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DEEL 106 AFLEVERING 9 | mmivroerrd963
TIJDSCHRIFT
VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
DE NEDERLANDSCHE ENTOMOLOGISCHE VEREENIGING
INHOUD:
Titelpagina en register van deel 106
| Tijdschrift voor Entomologie, deel 106, afl. 9. Gepubliceerd 2-XII-1963 |
Nederlandsche Entomologische Vereeniging
BESTUUR
Prof. Dr. J. van der Vecht, President (1961—1967), Oegstgeest.
Dr. J. G. Betrem, Vice-President (1959—1965), Deventer.
W. Hellinga, Secretaris (1963—1969), Amsterdam.
Drs. H. Wiering, Penningmeester (1962—1968), Bergen (N.H.).
Drs. C. A. W. Jeekel, Bibliothecaris (1960—1966), Amsterdam.
F.C. J. Fischer (1958—1964), Rotterdam.
Dr. H. J. de Fluiter (1960—1965), Wageningen.
COMMISSIE VAN REDACTIE VOOR DE PUBLICATIES
Prof. Dr. J. van der Vecht (1961—1967), Oegstgeest.
P. Chrysanthus (1961—1964), Oosterhout (N.B.).
Dr. A. Diakonoff (1961—1964), Leiderdorp.
G. L. van Eyndhoven (1963—1966), Haarlem.
Dr. L. G. E. Kalshoven (1961—1964), Blaricum.
Prof. Dr. D. J. Kuenen (benoemd 1957), Leiden.
Dr. P. A. van der Laan (benoemd 1957), Bennekom.
B. J. Lempke (1962—1965), Amsterdam.
Prof. Dr. J. de Wilde (benoemd 1957), Wageningen.
Dr. J. T. Wiebes (1963—1966), Leiden.
BESTUUR DER AFDELING VOOR TOEGEPASTE ENTOMOLOGIE
Dr. H. J. de Fluiter, Voorzitter, Wageningen.
Dr. F. J. Oppenoorth, Secretaris, Utrecht.
Mej. Dr. A. Post, Wilhelminadorp.
Dr. C. F. A. Bruyning, Oegstgeest.
‘Ir. P. Gruys, Rheden.
De contributie voor het lidmaatschap bedraagt f 15.—, voor student-leden
f 2.50, per jaar. — Begunstigers betalen jaarlijks tenminste f 15.—.
De leden, behalve de student-leden, ontvangen gratis de Entomologische
Berichten van 12 nummers per jaar, waarvan de prijs voor student-leden f 1.50
per jaar, voor niet-leden f 20 — per jaar en f 2.— per nummer bedraagt.
De leden kunnen zich voor f 10 — per jaar abonneren op het Tijdschrift voor
Entomologie; hiervan bedraagt de prijs voor niet-leden f 35 — per jaar.
De eerder verschenen publicaties der Vereeniging zijn voor de leden tegen ver-
minderde prijzen verkrijgbaar.
TIJDSCHRIFT VOOR ENTOMOLOGIE
Het Tijdschrift voor Entomologie wordt uitgegeven door de Nederlandsche
Entomologische Vereeniging en is bestemd voor de publicatie van de resultaten
van de studie der Entomologie van algemene en bijzondere aard. Het verschijnt
in één deel van 300—400 bladzijden per jaar, bestaande uit enkele afleveringen.
Bovendien worden monographieën handelende over bijzondere onderwerpen,
met onregelmatige tussenpozen uitgegeven.
DEEL 106 1963
TIJDSCHRIFT
VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
DE NEDERLANDSCHE ENTOMOLOGISCHE VEREENIGING
|
|
Tijdschrift vodk EME deel 106, 1963
Aflevering 1 verscheen 1 mei 1963
Afleveringen 2—5 verschenen 15 mei 1963
Aflevering 6 verscheen 13 augustus 1963
Aflevering 7 verscheen 16 september 1963
Afleveringen 8—9 verschenen 2 december 1963
REGISTER VAN DEEL 106
* Eén sterretje duidt een naam aan nieuw voor de wetenschap
* An asterisc denotes a name new to science
** Twee sterretjes geven aan een soort of vorm nieuw voor de Nederlandse fauna
** Two asteriscs denote a species or form new to the Netherlands fauna
The names of genera and species of the following groups which are listed at the end of
the respective papers, are not repeated in the present index. For these lists the reader is
referred to the following pages of the present volume:
Agaonidae (Hymenoptera), p. 109
Their fig hosts (Ficus species, Plantae), p. 110
Myriopoda, p. 205
Bactra species, etc. (Lepidoptera, Tortricidae), p. 356.
DIPLOPODA
Diplopoda 205
Paradoxosomatidae 205
Polydesmida 205
DIPTERA
Agromyza(idae, inae)
[113, 116
airae ll, 123
albipennis 116, 117,
[119—121, 122, 124, 161
ambigua 114—116
*baetica 118—120, 122,
12392152
*brachypodii (ssp.) 120,
122, 724, 75)
catherinae 116
celtidis 116
cinerascens 116
distorta 114, 118—121,
[123, 158, 161
*fennica (ssp.) 122, 128
frontosa 116
graminicola 118—120,
[122, 123, 148
graminivora 116
*hendeli 116—120, 122,
R 23s 16
intermittens 116
lucida 117, 119, 120, 123,
[133, 161
lunulata 121, 160
luteitarsis 116
megalopis 116
mobilis 116
nigrifemur 116
nigripes 113—121, 124,
[128, 161
nigrociliata 116
niveipennis 116
ocellaris 116
oliviae 116
oryzae 114, 118—121,
[123, 153
pallidifrons 116
parvicornis 116, 119
penniseti 116
phragmitidis 116—120,
1222123, 143
Poëmyza 161
prespana 119—123, 146
rubi 116
“spenceri 116, 118—120,
[123, 141
susannae 116
veris 116
yanonis 116
HYMENOPTERA
Agaonidae 1
Antrusa 161, 162, 164
Braconidae 161
Ceratosolen 1
Chalcididae 161
cincta 162, 164
Dacnusa 162—164
Dacnusini 161
Eulophidae 161
lugubris 163
maculata 184
melanocera 161, 164
ninella 164
nydia 162, 164
Opius 161
pione 162, 164
Vespa 184
LEPIDOPTERA
Acraea 197
affinis 197
alimena 199
Anaphaeis 197
andromacha 197
Appias 197
Bactra 285
Badamia 197
bolina 198, 199
cardui 197
Catopsilia 197, 199
chrysippus 197
corinna 198
crocale 197
Danaidae 198
Danaus 197—199
Delias 197
ega (var.) 197
eichhorni 198
Euploea 198
exclamationis 197
hamata (Danaus)
[197—199
hamata (Euploea) 198
harpalyce 197
Hypolimnas 198, 199
java 197
kershawi (var.) 197
lamina (var.) 199
nerina (var.) 198, 199
nigrina 197
Nymphalidae 199
paulina 197
petilia (var.) 197
plexippus 197
pomona 197, 199
pytanthe 197
pythias (var.) 197
446
septentrionis (var.) 199
smilax 197
Terias 197
teutonia 197
Tortricidae 285
Vanessa 197
ORTHOPTERA
albomarginatus 358, 375
biguttulus 358, 374
brunneus 374
Chorthippus 358, 374, 375
coerulescens 375
lineatus 384
maculatus 358, 374
mollis 375
Myrmeleotettix 358, 374
Oedipoda 375
Omocestus 375
parallelus 375
Stenobothrus 384
viridulus 375
RHYNCHOTA
Aepophilus 169, 170, 174,
[178, 189
Amphibicorisae 170
Berytidae 186
bonnairei 169, 170, 174,
[178, 189, 193
bouchervillei 186
Chiloxanthus 188
Cicicomorpha 170, 189
Dysdercus 182, 184
Gelastocoris 181, 182
Gerridae 186
Heteroptera 169
Homoptera 182
Hydrocorisae 170, 189
intermedius 182, 184
littoralis 188
Lygaeidae 190
Mesovelia 169—172, 174,
[178, 180, 188, 189, 193
Mesoveliidae 169, 189
Miespa 186
muelleri 186
mulsanti 169, 171, 174,
[178, 188, 190, 193
Nabidae 186
Naucoris 184
Notonecta 184
pallipes 186
palustris 169, 170, 176,
[180, 193
Pentatomomorpha 170,
[189
pilosella 185
pilosus 188
Salda 186, 188
Saldidae 169, 189
Saldoida 184, 186
Saldula 169—171, 174,
[176, 180, 181,
[184, 189, 193
saltatoria 182
splendida 186
vittigera 170, 172, 175,
[180, 190, 193
THYSANOPTERA
Aeolothrips 201, 202
albopictus 204
alni 203
Anaphothripina 202
annulipes 203
Aptinothrips 202
bidens 203
Cephalothrips 204
consociatus 201
Cryptothrips 203
#*cytisi 201, 202
elegans 202
ericae 201
Halothrips 203
Hoplandrothrips 203
Hoplothrips 204
Iridothrips 202
juncorum 201
kratochvili 202
longisetis 204
loti 201
manteli 202
mariae 202
meliloti 202
minutissimus 203
monilicornis 204
muelleri (f.) 201
Neoheegeria 203
nigripes 203
obscura (f.) 203
obscurus 201
Odontothrips 201, 202
phaleratus 202
Phlaeothrips 203
Poecilothrips 204
propinquus 202
Rhopalandrothrips 201
subterraneus 203
Taeniothrips 202
Thrips 203
Tubulifera 203
ulicis 201
ulmi 204
verbasci 203
williamsianus 203
Xaloplothrips 203
PLANTAE
Agrosteae 120
Agrostis canina 120
A. stolonifera 120
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 9, 1963
Alnus incana 203
Anchusa officinalis 202,
{203
Arrhenatherum elatius
[120
Arundineae 119, 120
Aveneae 120
Brachypodieae 120
Brachypodium sylvaticum
1201220194
Cerastium arvense 201
Deschampsia caespitosa
[119, 120, 123
Echinaria 120
Echium vulgare 202
Fagus sylvatica 204
Festuca(ceae) 120
Ficus 1
Genista pilosa 201
Glyceria(eae) 119, 120,
[122—124
G. fluitans 120
G. maxima 120, 121
Graminaceae 113, 116,
[119, 202—204
Holcus 120, 122, 124
Hordeae 119—121
Hordeum 120, 121
H. vulgare 127
H. murinum 127
Lilium 203
Malus 203
Melilotus officinalis 202
Oryza sativa 120, 121, 123
Paniceae 120
Pennisetum 116
Phalaris 119, 122, 124
P. arundinacea 120
P. erucaeformis 120
Phleum pratense 120
Phragmites communis
ye) 102, 125
Poa 120
Prunus avium 204
Quercus 204
Reseda lutea 201
Sarothamnus scoparius
[201, 202
S. vulgaris 202
Secale cereale 120
Seslerieae 120
Setaria viridis 120
Sorbus aucuparia 202
Symphoricarpus rivularis
[203
Triticum 121
id. aestivum) 120, 122,123
Typha latifolia 202
Ulmus carpinifolia 204
Verbascum 202, 203
Vicia cracca 201
Zea mays 116
Zizonia latifolia 120, 121,
[123
2 Ae
TIME
(TA
ee
N
Mi
wait
INHOUD VAN DEEL 106
DIAKONOFF, A. — African species of the genus Bactra Stephens (Lepido-
ptera, Tortricidae) ee Lee: :
FRANSSEN, C. J. H. & W. P. MANTEL — Supplement op de Naamlijst van
de in Nederland aangetroffen Thysanoptera . SP RATIO A
GRIFFITHS, G. C. D. — A revision of the Palaearctic species of the nigripes
group of the genus Agromyza Fallén (Diptera, Agromyzidae) .
Gupta, A. G. — Comparative dai of the Saldidae and Meso-
veliidae (Heteroptera) . PROTEINA
JEEKEL, C. A. W. — Paradoxosomatidae from Borneo (Diplopoda,
Polydesmida) BEN RN
LENSINK, B. M. — Distributional ecology of some Acrididae (Orthoptera)
in the dunes of Voorne, Netherlands . ARA eN,
MANTEL, W. P. — zie FRANSSEN & MANTEL.
STRAATMAN, R. — Observations on migration in certain Australian Lepido-
ptera .
WIEBES, J. T. — Taxonomy and host preferences of Indo-Australian fig
wasps of the genus Ceratosolen (Agaonidae) .
Register van deel 106.
NO)
O0
WN
No
S)
Ww
357
Hol ' or
eat ad A
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BD —1 568.2 Tuus. var. 2001 |
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Ip}!
DEEL 106 AFLEVERING 1 | HARVARD 1943
| i UNIVERSITY |
A da
TIJDSCHRIFT
VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
DE NEDERLANDSCHE ENTOMOLOGISCHE VEREENIGING
INHOUD:
J. T. WieBEs. — Taxonomy and host preferences of Indo-Australian fig wasps of
the genus Ceratosolen (Agaonidae), pp. 1—112, figs. 1—329, pl. 1.
Tijdschrift voor Entomologie, deel 106, afl. 1. Gepubliceerd 1-V-1963 |
Nederlandsche Entomologische Vereeniging
BESTUUR
Prof. Dr. J. van der Vecht, President (1961—1967), Oegstgeest.
Dr. J. G. Betrem, Vice-President (1959— 1965), Deventer.
W. Hellinga, Secretaris (1963—1969), Amsterdam.
Drs. H. Wiering, Penningmeester (1962—1968), Bergen (N.H.).
Drs. C. A. W. Jeekel, Bibliothecaris (1960—1966), Amsterdam.
F. C. J. Fischer (1958—1964), Rotterdam.
Dr. H. J. de Fluiter (1960—1965), Wageningen.
COMMISSIE VAN REDACTIE VOOR DE PUBLICATIES
Prof. Dr. J. van der Vecht (1961—1967), Oegstgeest.
P. Chrysanthus (1961—1964), Oosterhout (N.B.).
Dr. A. Diakonoff (1961—1964), Leiderdorp.
G. L. van Eyndhoven (1963—1966), Haarlem.
Dr. L. G. E. Kalshoven (1961—1964), Blaricum. : |
Prof. Dr. D. J. Kuenen (benoemd 1957), Leiden. |
Dr. P. A. van der Laan (benoemd 1957), Bennekom.
B. J. Lempke (1962—1965), Amsterdam.
Prof. Dr. J. de Wilde (benoemd 1957), Wageningen.
Drs. J. T. Wiebes (1963—1966), Leiden.
BESTUUR DER AFDELING VOOR TOEGEPASTE ENTOMOLOGIE
Dr. H. J. de Fluiter, Voorzitter, Wageningen.
Dr. F. J. Oppenoorth, Secretaris, Utrecht.
Mej. Dr. A. Post, Wilhelminadorp.
Dr. C. F. A. Bruyning, Oegstgeest.
Ir. P. Gruys, Rheden.
De contributie voor het lidmaatschap bedraagt f 15.—, voor student-leden
f 2.50, per jaar. — Begunstigers betalen jaarlijks tenminste f 15.—.
De leden, behalve de student-leden, ontvangen gratis de Entomologische
Berichten van 12 nummers per jaar, waarvan de prijs voor student-leden f 1. 50
per jaar, voor niet-leden f 20.— per jaar en f 2.— per nummer bedraagt.
De leden kunnen zich voor f 10.— per jaar abonneren op het Tijdschrift voor
Entomologie; hiervan bedraagt de prijs voor niet-leden f 35.— per jaar.
De eerder verschenen publicaties der Vereeniging zijn voor de leden tegen ver-
minderde prijzen verkrijgbaar.
TIJDSCHRIFT VOOR ENTOMOLOGIE |
Het Tijdschrift voor Entomologie wordt uitgegeven door de Nederlandsche |
Entomologische Vereeniging en is bestemd voor de publicatie van de resultaten |
van de studie der Entomologie van algemene en bijzondere aard. Het verschijnt |
in één deel van 300—400 bladzijden per jaar, bestaande uit enkele afleveringen.
Bovendien worden monographieën handelende over bijzondere onderwerpen,
met onregelmatige tussenpozen uitgegeven.
oy
um
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963 PLAAT 1
wen mann
2, oh ee BE id
Fee De arani
Pl. 1, Ceratosolen vechti Wiebes, female and male, X 30. H. HEIJN del.
J. T. WIEBES : Indo-Australian Ceratosolen
| Niens:
TAXONOMY AND HOST PREFERENCES “INDO-
AUSTRALIAN FIG WASPS OF THE GENUS CERATOSOLEN
(AGAONIDAE)
BY
J. T. WIEBES
Rijksmuseum van Natuurlijke Historie, Leiden
CONTENTS
odio So LIO CR RER Se ee 1
The Indo-Malayan and Papuan species of the genus Ceratosolen … nnen 3
kaxonomiEhteviston BAE EN TTI dede: 3
Glassificattonmwancikevastomthers Decies ene 84
iemhostmpreterencesWofntheimAoaonid ach 92
Speciticity of the relation between fig) wasps and) Fiss... 92
Bhylogenetteanspeciticity ne A OO NATO TE I 99
Concludinemrenmatks MIE ANNAE RI 105
DE CIENCCS MME ee een EEE 106
oee TREE en OL 109
INTRODUCTION
The systematic position of the fig wasps has for a long time been a challenging
problem to biologists. Up to about 1880, the literature on fig wasps consisted of
only a few occasional papers on the curious creatures found in the receptacles of
Ficus species, and on the function they were supposed to fulfil, or were denied
to have, in the process of caprification. For a survey of the literature prior to
1880, I refer to the critical review by MAYER (1882).
SAUNDERS (from 1878 onwards) and WESTWOOD (1882, 1883) described a
number of genera and species of fig wasps from India and Australia.
PAUL MAYER (1882) reported upon a collection of fig insects from several parts
of the world, mainly collected by H. zu SOLMS-LAUBACH in the Botanical Gardens
at Bogor (Java), and by F. MULLER in Brazil.
Gusrav Mayr (1885) described and named the species listed by MAYER and
some additional material sent by SoLMs from Bogor, thus providing a firm basis
for the study of the fig wasps. In 1906, MAYR contributed an additional paper,
consisting of corrections to his 1885 paper and descriptions of new species and
genera.
In the meantime ASHMEAD (1904) reviewed the classification of the Chalcid-
oidea, and assigned the fig wasps to their proper place in this large group of
Hymenoptera.
From 1916 onwards, the knowledge of the fig wasps was greatly enlarged by
GRANDI, who published some fifty-five papers on their taxonomy, morphology,
1
2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
and bionomics. A survey of GRANDIs work may be found in GRANDI (1955c,
1961). Among his outstanding contributions to the knowledge of the Agaonidae,
the critical review (1928c) of the species described by Mayr, the monograph of
Blastophaga psenes (second edition, 1929), and the world catalogue (last edition, |
1955b) especially may be mentioned.
Up to a few years ago, only little attention could be given to the host relations
of the Agaonidae, due to the fact that it was very difficult to have the host Ficus
properly identified. Although MAYR (1885) cooperated with the botanist SOLMS,
and WILLIAMS (1921, 1928) with MERRILL, the host preferences could in many
instances not be ascertained.
It was a fortunate coincidence that at the time when Dr. J. VAN DER VECHT
took up the study of fig wasps in the Botanical Gardens at Bogor, Java, Mr. E. J.
H. CORNER was working on a revision of the genus Ficus. They cooperated in the
study of the symbiosis between figs and fig wasps, and pointed out the fact that
the relation between figs and fig wasps is highly specific. VAN DER VECHT (1956,
p. 103; 1960) emphasized the particular importance of the study of the symbio-
sis, “for probably nowhere will botanists and entomologists find such an in-
teresting opportunity to compare and check their ideas about certain problems of
natural relationships and evolution”.
VAN DER VECHT started his studies on fig wasps in 1954, and collected many
samples in Java. In 1955, however, he returned to Holland, and soon found him-
self engaged in other work. In 1959 he suggested that I should proceed with the
subject, and kindly gave the collection and his field notes in my care.
In the meantime CORNER enlarged the collection with many samples of wasps
taken from figs preserved in botanical collections. A few years later he sent to
me two collections of fig insects made on his expeditions to New Guinea, Melanesia,
and Borneo. This material, together with the collections made by VAN DER VECHT,
is preserved in the Rijksmuseum van Natuurlijke Historie, Leiden.
It soon became evident that the outcome of my study of the fig wasps could only :
gain by the incorporation of the data of collections available from other museums. —
Upon my request, material was sent on loan by various institutions. The material
on which the present paper is based belongs to the institutions mentioned below;
the names are preceded by the abbreviations used in the text of this paper.
BM — British Museum (Natural History), London;
BMH — Bernice P. Bishop Museum, Honolulu;
HSPA — Hawaiian Sugar Planters’ Association, Honolulu (the collection of DE
F. X. WILLIAMS);
ML — Rijksmuseum van Natuurlijke Historie, Leiden (in most instances,
however, indicated by a collection number only);
NMB — Naturhistorisches Museum, Basel;
NMW — Naturhistorisches Museum, Wien;
OUM — Hope Department of Entomology, University Museum, Oxford;
USNM — United States National Museum, Washington.
. . . . . |
Moreover, some typical specimens were received in exchange for specimens from
the collection ML from Dr. G. GRANDI (Bologna) and Dr. K. J. JOSEPH (Dia
war, India) or in loan from Dr. E. F. RıEK (Canberra).
J. T. WieBEs: Indo-Australian Ceratosolen 3
Because of practical reasons I restrict the work to the Indo-Malayan and Papuan
areas. The bulk of the material is from Malaya, Indonesia, and New Guinea, and
these are the areas in which CORNER made an extensive research on the genus
Ficus. As Dr. E. F. RIEK informed me that he has prepared a manuscript on the
Australian Chalcidoidea, I do not include the Australian species. Only some
Australian records of Indo-Malayan species are mentioned.
All information on Frcus mentioned in the present paper was either given by
CORNER (personal communication, and in litt. to VAN DER VECHT and WIEBES),
or taken from his publications (CORNER, 1958, 1960—1962).
Acknowledgements.
For loan of material, and for generous help in numerous ways, I am indebted to
the institutions mentioned above and to the following persons: J. H. ARDLEY
(Lae), F. A. BIANCHI (Honolulu), B. D. BurKs (Washington), J. H. CALABY
(Canberra), M. FISCHER (Vienna), M. DE V. GRAHAM (Oxford), G. GRANDI
(Bologna), K. J. JosePH (Dharwar), F. Keiser (Basel), J. F. PERKINS (London),
E. F. Riek (Canberra), Miss. M. T. WieBEs (Leiden), and C. M. YOSHIMOTO
(Honolulu).
The work on this paper could not have begun without the collection of
VAN DER VECHT as a firm working basis, and it could not have been completed
without the continuous cooperation of Mr. E. J. H. CORNER. I am grateful to Dr.
J. VAN DER VECHT for entrusting me with this research, and for valuable com-
ment and advice. To be permitted to cooperate with Mr. CORNER is a highly
esteemed privilege.
A grant from the “Zoölogisch Insulinde Fonds” enabled me to travel to Cam-
bridge, Oxford, and London, in order to discuss some problems with Mr. CORNER,
and to go through the collections of the Oxford University Museum (Hope
Department of Entomology) and the British Museum (Natural History).
The Nederlandsche Entomologische Vereeniging kindly accepted this paper for
publication.
THE INDO-MALAYAN AND PAPUAN SPECIES OF THE GENUS
CERATOSOLEN
TAXONOMIC REVISION
Ceratosolen Mayr
Sycocrypta Coquerel, 1855, pp. 367, 422 (monobasic, type: S. coeca Coquerel); Walker,
1871, p. 60 (in family Agaonidae s.1.); Newman, 1871, p. 400 (do.); Saunders, 1878, pp.
316—317 (in synonymy of Blastophaga Gravenhorst); Westwood, 1882, p. 48 (do.);
Mayer, 1882, p. 586 (do.); Westwood, 1883, p. 379 (do.); Mayr, 1885, pp. 187—188
(incertae sedis); Dalla Torre, 1898, p. 321 (catalogue); Ashmead, 1904, p. 389 (in syno-
nymy of Blastophaga Gravenhorst); Gahan & Fagan, 1923, p. 138 (types of Chalcid genera);
Wiebes, 1961a, p. 236 (synonymous with Ceratosolen Mayr); 1961c, pp. 283—384 (proposed
suppression).
Ceratosolen Mayr, 1885, pp. 150, 159, 160, 162, 164 (seven species; subgenus of Blas-
tophaga Gravenhorst, key, description, key to species); Dalla Torre, 1898, pp. 323—325
(catalogue); Mayr, 1906, p. 153 (treated as separate genus); Schmiedeknecht, 1909, pp. 7,
9—10 (key, catalogue); Grandi, 1916a, pp. 133—153 (description, key to species); Gahan &
Fagan, 1923, p. 29 (types of Chalcid genera); Aiyar, 1925, pp. 236—237 (catalogue of
4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Indo-Ceylonese species); Grandi, 1928c, pp. 221—225 (catalogue); 1935, pp. 225—229
(do.); Mani, 1938, pp. 29—30 (catalogue of Indian species); Grandi, 1941, pp. 13—18
(catalogue); 1952c, pp. 81—85 (do.); 1955b, pp. 120—125 (do.); Wiebes, 1961a, pp.
236, 238 (synonymous with Sycocrypta Coquerel, key); 1961c, pp. 383—384 (proposed
validation).
Ceratosolens {!} Ashmead, 1904, pp. 233, 234 (key, type designated: Blastophaga (Cera-
tosolen) appendiculata Mayr).
Ceratosolensia Girault!), 1915, p. 311 (monobasic, type, originally designated: C. ficophaga
Girault); Gahan & Fagan, 1923, p. 29 (types of Chalcid genera); Grandi, 1928c, p. 229
(catalogue, incertae sedis); 1935, p. 239 (do.); 1941, p. 27 (do.); 1952c, p. 94 (do.);
1955b, p. 135 (do.).
Restriction.
As may be seen from the synonymy, the genus Ceratosolen was originally estab-
lished as a subgenus of Blastophaga Gravenhorst. The most important differential
characters are the situation of the male antennae in separate grooves, and the
elongate shape of the spiracular peritremata of the eighth urotergite in the female.
In 1906 Mayr, without explicitly stating the change, treated the group as a
separate genus, an opinion adhered to by all subsequent authors.
It appears, however, that there are some species that constitute a grade between
Blastophaga and Ceratosolen. Especially in the figs of the section Sycidium Miq.,
I find species of wasps that show Blastophaga characters in the male, whereas the
female has large, elongate, instead of circular peritremata on the eighth abdominal
segment; other species have females with circular peritremata, and males with a
Ceratosolen facies. Suffice it here to mention the gradation in these few characters
only.
The genus evidently needs redefinition against Blastophaga and against some
genera named by GIRAULT. This redefinition, however, cannot be given before
the other genera have been revised. In this paper, the genus Ceratosolen is treated in
its restricted sense, excluding the species that do not show the above-mentioned
differential characters in both sexes.
IDESERI PROM
For an extensive diagnosis of the genus, I refer to GRANDI (1916a, pp. 133—
153). The following description may serve to facilitate recognition, and to elu-
cidate some terms and notations used in this paper.
The quotients given in parentheses with the descriptions of some body-parts —
e.g., pronotum (10:7) — refer to the relative length and width, respectively; if
given in square brackets behind a comparative remark — e.g., femur longer than
tibia [10 : 7} — they refer to the comparison.
The morphological terms used in the descriptions are the English equivalents of
those used by GRANDI (1929) in his monograph on Blastophaga psenes (L.). In
1) Dr. E. F. RIEK, who studied the type specimens of GIRAULT's Australian Chalcodoidea, |
drew my attention to this new synonym, and kindly sent to me some female specimens of |
C. ficophaga for comparison with the other species of Ceratosolen. C. ficophaga seems to |
belong to the group of C. appendiculatus (Mayr), but the males should be studied for a |
definitive classification of the species.
J. T. WIEBES : Indo-Australian Ceratosolen 5
a few cases, e.g., the venation of the wings, I use different names, which are
self-evident.
Male. Head longer than wide. The head may be pubescent, or provided with
small spines. In most cases, these adornments are omitted from the figures.
Epistomal margin usually trilobate, bilobate in one species; always prominent and
distinctly visible. Antenna — not counting the radicola — consisting of four to
six segments: scape, pedicel, flagellar segments; not clubbed. Unless otherwise
stated, the lengths of the segments are measured along the dorsal edges. In
preparing the figures, care was taken not to press the antenna between slide and
cover-glass. Contrarily, the mouthparts were flattened into one plane, so that they
may look different when studied in situ. Labium and maxillae sometimes atro-
phied, mostly present; maxillae with or without distinct lateral expansions. Mandi-
ble bidentate, each tooth with a subapical expansion.
Thorax simple; dorsally, the following terga are visible: pronotum, mesonotum,
metanotum. The metanotum is, in most species, incompletely separated from the
propodeum. Propodeum, and its spiracular peritremata, variable in shape. Ven-
trally, the following sterna are visible: prosternum (with the propleurae indicated
in the figures by a dotted line), mesosternum, and metasternum. The metasternum
is hirsute in some species.
Tibia of the fore leg with a varying number of apical teeth. Tarsus consisting of
two or three segments, which are sometimes incompletely separated. Tibiae of mid
and hind legs with or without spines on the disks. The normal number of tarsal
segments is five, although in one species there are only three tarsal segments
in the mid and hind legs, and in other species oligomery or heteromery
occurs. Unless otherwise stated, the length of the distal tarsal segment is measured
without the claws. In many species the hind legs are hirsute, and the tarsi are
greatly dilated. The figures of these parts are semi-diagrammatical, merely in-
dicating the size of the segments and the length of the pubescence.
Gaster. The tenth urite may or may not bear cerci or parameres, or both; the
aedeagus is in most instances dilated subapically.
Length. The measurements given refer to the total length of head, thorax, and
propodeum. The colour is that of specimens preserved in alcohol.
Female. Head approximately as long as wide across the compound eyes. The
length of the cheek, given in comparison with the longitudinal diameter of the
eye, refers to the distance from the lower margin of the eye to the point of in-
sertion of the mandible. Three ocelli. Lateral expansions of the epistomal margin
more or less angular, or rounded. Antenna relatively simple, the club in most
species only indicated by the shape of the loosely connected apical two or three
segments, or absent; in other species the apical two or three segments completely
united. Maxilla with or without a bacilliform process. Mandible, and its appen-
dage, with a varying number of ventral ridges.
Thorax. Wings pubescent, or with glabrous parts. Venation of the fore wing
complete, i.e., consisting of submarginal, marginal, stigmal, and postmarginal
veins. Stigma sometimes with dark radiating striae.
Fore leg with a distinct comb on the dorso-apical edge of the tibia, which may
6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
consist of a varying number of teeth. In some cases, the ventral tooth of this comb
is blunt. Ventro-apical edge with smaller teeth, and, mostly, with a long, simple
spur. Tarsus usually pentamerous, in a few species consisting of only four seg-
ments. Mid leg slender; tarsus pentamerous, tetramerous in two species. Hind tibia
with apical teeth, which may vary in shape and number. Hind tarsus pentamerous.
Gaster. The spiracular peritremata of the eighth urotergite elongate, gouge-like.
The ovipositor projecting beyond the gaster to a varying extent.
Length. The measurements given refer to the total length of head, thorax, and
gaster (without the ovipositor). Colour as seen in the alcohol samples.
Bionomics.
Two authors gave notes on the bionomics of species of Ceratosolen, viz., BAKER
(1913) on C. notus (Baker), and WILLIAMS (1928) on the same species, on
C. appendiculatus (Mayr) (not indicated by name), and on C. bakeri Grandi.
The following short description of the life of a species of Ceratosolen — nearly
all species of which live in dioecious figs — is almost verbally borrowed from
WILLIAMS (1928, p. 7).
The wasps issue from the gall flowers of a mature fig. The males, which are
quite flightless, hatch first and immediately seek the females by biting open the
occupied galls. After mating the males perish, with some exceptions within the
receptacles. On the other hand the females are fully winged, and in seeking egress
from the receptacle through the ostiole or by a hole drilled through the rind, in-
advertently dust themselves with pollen from the ripe stamens. They take wing,
and under proper conditions find and enter a young fig in the right stage of
development, i.e., one whose stamens are as yet undeveloped but whose gall
flowers and seed flowers are in the one instance ready and adapted to receive the
egg, and in the other instance, the pollen to fertilize the seed-producing ovary.
Evidently, the females are unable to distinguish between the male and the female
fruits, and enter whatever sort is available. Of course, the wasp cannot reproduce
in the seed or female fig because this is not modified to that end, neither does it
find fig flowers to fertilize in the male or gall fig.
The young fig grows to maturity, the stamens and seeds ripen, and wasps issue
from the galls.
Remarks.
Under each species I give a list of synonyms — not necessarily exhaustive, as
the references to the catalogues are not included — the data on the material stud-
ied, a description or descriptive notes, if necessary, and, in most cases, general
remarks.
In general, the geographical names mentioned are those now in common use
(e.g., Bogor, not Buitenzorg).
Hosts identified by CORNER are indicated as such, other host names are given as
I found them on the labels. Discussion of the host records may be found in the
chapter on host preferences.
|
J. T. WIEBES : Indo-Australian Ceratosolen 7
Fig. 1, Ceratosolen hewitti, Singapore, male, 2—5, C. constrictus, Java, 2, male, 3, female
fore tibia, 4, female hind leg, 5, apex of male abdomen, 6, C. pygmaeus, Mindanao, apex of
male abdomen, 7—13, C. marshalli, Fiji, male, 7, mandible, 8, antenna, 9, cercus of tenth
urite, 10, male, 11, fore leg, 12, hind leg, 13, mid leg. Figs. 1, 2, 10, X 50; 4, X 80;
Use, NS B L'ENCRE GL SCI
8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Katozusgieis.
If not otherwise stated, the following body-parts are figured as seen in dorsal
aspect: male head and thorax, antenna; figured as seen in ventral aspect: male and
female mouthparts, male genitalia; as seen in antaxial aspect: male and female
legs (i.e., retrolateral aspect of fore leg, prolateral aspects of mid and hind legs),
female antenna.
As there are some differences between the various samples in the state of
preservation, all specimens from coll. HSPA and BMH were sent to me dry,
and they had to be relaxed in alcohol, not all figures of comparable body-parts
could be magnified to the same, large scale.
But for the plate, the figures are original camera lucida drawings by the
author.
Ceratosolen pygmaeus Grandi (fig. 6)
Ceratosolen pygmaeus Grandi, 1927a, pp. 317—320, Pl. 4 figs. 50—65, Pl. 5 fig. 66
[descr. 9, &, ex Ficus minahassae Miq., Mt. Maquiling (Luzon, Philippine Is.), leg. C.
F. Baker}.
Mirae rican:
One 9, 1 4, ex Ficus minahassae Miq., Luzon (Philippine Is.), ex coll.
GRANDI; coll. ML, no. 508.
Series 9,1 4, ex Ficus minahassae Mig., Los Baños (Luzon, Philippine Is.),
Goll Agric lege Fy Xe WAE MSIE NAO 21 coll ’SEISPAr
Series 9, 4, ex Ficus minahassae Mig., Lake Lanao (Mindanao, Philippine
IS) Me ME AV Erra is XT.192T coll EISPA CON MIE no 5 SSM PONG
Description — Additional note.
Male. Labium and maxillae completely atrophied. The tenth urite bears cerci,
which are, due to the smallness of the specimens, and to the fact that these ap-
pendages are almost hyaline, very difficult to be observed. In the specimens from
the HSPA collection, the cercus bears one claw (fig. 6).
Female. Tarsi of the fore and mid legs heteromerous.
Ceratosolen marshalli Grandi (figs. 7—13)
Ceratosolen Marshalli Grandi, 1931, pp. 8—11, figs. III [descr. ®, ex Ficus spec.,
Mt. Labasa (Fiji Is.), leg. W. Greenwood, 1.VII.1923, no. W. G. 532!) }.
Material.
Series 9, &, ex Ficus spec. near storcki, Fiji, leg. C. E. PEMBERTON, 1920;
coll. HSPA, 1 & (allotype) slide mounted; coll. ML, no. 595: 10 9,2 4; 8,
slide 595a, 9, 595b, c.
Description — Additional note.
Female. The specimens from the collection WILLIAMS are very similar to the
1) CORNER studied W. G. 532, and identified it with Ficus pritchardii Seem.
J. T. WIEBES : Indo-Australian Ceratosolen 9
species described by GRANDI as C. Marshall. There are some minor differences,
viz., in the specimens seen by me, the mandibular appendages bear five ventral
ridges instead of four, and the sensilla of the funicular segments seem to be more
numerous. As the material has been desiccated, as have GRANDI's specimens, the
last mentioned character is not easily to be observed. The colour-differences
between head, thorax, and gaster, mentioned by GRANDI (Lc, p. 8): ‘colore
fondamentale isabellino-ferrugineo sfumato di umbrino, con la parte posteriore del
cranio e gli urotergiti tendenti al fuligineo’’, are very conspicuous. I consider the
sample conspecific with GRANDI's species, and describe the hitherto unknown
male.
Male. Head (fig. 10) not quite twice as long as wide. Lateral lobes of the
epistomal margin prominent, median lobe small. Eyes absent. Antennal grooves
half closed. Antenna (fig. 8) four-segmented: scape (8 : 5) slightly longer than
the pedicel (3 :2), which is two-thirds the length of the next segment (9 : 4),
Third segment nearly as long as the apical segment (3 : 1). Mandible, fig. 7;
labium and maxillae atrophied.
Thorax, fig. 10. Length of the pronotum four-thirds of the posterior width, and
nearly twice the anterior width. Mesonotum twice as wide as long. Metanotum
incompletely separated from the propodeum, approximately as wide as the meso-
notum. Propodeum wider than long {4 : 3}, narrower than the mesonotum. Fore
leg (fig. 11): femur more than twice as long as the tibia, which bears three dorsal
and two ventro-apical teeth. Tarsus bimerous, segments in ratio 5 : 6. Mid leg
(fig. 13): the coxa two-thirds the length of the clavate femur, the trochanter
small. Tibia arcuate, as long as femur and trochanter combined, with acute apical
edges. Tarsus pentamerous, the segments in ratio 8: 3: 3: 4: 9. Femur of the hind
leg (fig. 12) slightly larger than the coxa; the tibia about as long as the femur,
with three apical teeth. Tarsus pentamerous, the segments in ratio 8:3:3:4:8.
Gaster. Cerci (fig. 9) of the tenth urite with three rather robust claws.
Length, 1.1—1.3 mm. Colour light yellow-brown.
Remark.
There are several characters by which C. marshalli can be distinguished from
C. pygmaeus, but it is evident from other features (e.g., the dentation of the
fore legs, the constitution of the antennae) that the two are closely related.
Ceratosolen nanus sp. n. (figs. 14—25)
Material.
Series 9, 4, ex Ficus pungens Reinw. (det. E. J. H. CORNER), Lae (Terr.
New Guinea), leg. E. J. H. CORNER, 5.IX.1960; coll. no. 548; 4 (holotype),
slide 548a, @ (allotype), 548c, 4, 9 (paratypes), 548b, d.
DESC pito:
Male. Head (fig. 20) slightly more than twice as long as wide anteriorly, and
approximately one and a half times as long as its maximum width [13 : 9}. Dorsal
surface with very small bristles. Epistomal margin obtusely trilobate, the lateral
10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Figs. 14—25, Ceratosolen nanus, New Guinea, 14, male antenna, 15—18, female, 15,
mouthparts, 16, antenna, axial aspect, 17, fore leg, axial aspect, 18, antenna, detail, 19—23,
male, 19, mid leg, 20, male, 21, hind leg, 22, mandible, 23, detail of fore leg, 24, female
mid leg, 25, detail of female hind leg. Figs. 20, X 115; 17, 19, 21, 23—25, X 155; 1416,
US 22, SK AUS
J. T. WieBes: Indo-Australian Ceratosolen 11
lobes about twice as long as the median one, and without hairs. Eyes large. An-
tennal grooves half closed, obtuse behind. Antenna (fig. 14) five-segmented;
the scape (7 : 4) slightly longer than the pedicel (3 : 2), first flagellar segment
(7:6) half as long as the pedicel, second segment (1 : 1) somewhat longer than
the first, apical segment (3 :1) distinctly longer than the pedicel. Pedicel, first,
and second flagellar segments with one apical hair. Mandible, fig. 22; labium and
maxillae completely atrophied.
Thorax, fig. 20. Posterior width of the pronotum distinctly less than its length
[9 :11}. The pronotum is narrower in front than behind [7 : 9}. Length of the
mesonotum one half, that of the metanotum one third of the width. Propodeum
about one and a half times as wide as long {14:9}, with rounded edges. Tibia
of the fore leg (fig. 23) half as long as the femur, the tarsus (including the
claws) as long as the tibia. Tibia with two large teeth at the dorso-apical edge
and one small ventral tooth. Tarsal segments subequal, claws large. Mid leg
(fig. 19): coxa and femur subequal, tibia a little longer, without apical teeth.
Tarsus trimerous, the segments in ratio 4: 2:5. Coxa of the hind leg (fig. 21)
slightly longer than the femur; the tibia distinctly shorter, with ventral teeth.
Tarsus trimerous, the segments in ratio 5 :3 : 7.
Gaster. Genitalia without cerci.
Length, 0.95—1.0 mm. Colour yellowish.
Female. Head as long as wide across the eyes. Longitudinal diameter of the
compound eye longer than the cheek {7 :5}. Pubescence short. Antenna (figs.
16, 18): scape as long as fourth to seventh segments combined; pedicel not half as
long as the scape, with approximately twenty spines on the axial surface. Fourth
segment small. Fifth to eleventh segments gradually diminishing in length, the
ninth segment the widest. Segments with few sensilla: the fifth with three, the
sixth and seventh with five, the eighth with six, the ninth with seven, the tenth
with five, and the eleventh with two oblong sensilla; the ninth to eleventh seg-
ments with some circular sensilla. Mouthparts, fig. 15. Mandible with four ventral
ridges, longer than the appendage, which bears six ventral ridges. Labium and
maxillae much like those of C. pygmaeus.
Thorax. Pronotum broad and bristly. Scutum nearly as long as wide; scutellum
approximately as long as wide posteriorly, with a few scattered hairs. Metanotum
with many hairs. Propodeum as wide as scutum and scapulae combined, with tufts
of five hairs above and beneath the spiracular peritremata. Fore wing (16 :7),
1.1 mm long. Submarginal, marginal, stigmal, and postmarginal veins in ratio 8:
5 :4:2. Submarginal vein with three pustules, stigmal vein with four. Hind
wing (6:1), 0.6 mm long. Wings pubescent over nearly the whole surface,
marginal fringes long. Coxa and femur of the fore leg (fig. 17) subequal, the
tibia half as long as the femur. Dorsal apex of the tibia with two teeth. Tarsus
tetramerous, the segments in ratio 25 :14:12:15. Coxa of the mid leg (fig.
24) semi-globular, the tibia about as long as femur and trochanter combined.
Tarsus tetramerous, the segments in ratio 22 : 23:16:17. Hind leg (fig. 25):
the femur a little longer than the coxa, the tibia nearly as long as the coxa. Apical
tibial tooth bidentate. Tarsus pentamerous, the segments in ratio 10:6:5:3:4.
12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Gaster. Ovipositor barely projecting beyond the apex of the gaster.
Length, 1.2 mm. Colour light brown.
Remark.
This species is well characterized by the shape of the female antenna. The oligo-
mery of the legs and the general facies connect it with C. pygmaeus.
Ceratosolen constrictus (Mayr) and C. hewitti Waterston (figs. 1—5;
map, fig. 261)
It is possible that C. constrictus (Mayr) and C. hewitt: Waterston merely
represent two races of one species. C. constrictus is known from Java and Sumatra,
C. hewitti from Borneo and Malayat). Moreover, the specimens from Mt. Kina-
balu (N. Borneo) are provisionally referred to C. constrictus, For distribution,
see the map (fig. 261).
The males of C. hewitti differ from those of C. constrictus in the relative
proportions of head, thorax, and propodeum (figs. 1 and 2), and are slightly
longer. The females of C. hewitti have longer antennal segments than those of
C. constrictus, longer sensilla, and more hairs near the propodeal spiracle (cf. the
figures given by WATERSTON, 1921, and GRANDI, 1928c).
Although there are some characters connecting C. constrictus and C. hewitt with
C. crassitarsus and its relatives, the general facies, the heteromery of the male
mid and hind tarsi, and the absence of the tibial spur from the female fore leg,
are arguments for placing them in the relationship of C. pygmaeus.
Ceratosolen constrictus (Mayr) (figs. 2—5)
Blastophaga spec., Mayer, 1882, p. 571 [®, 4, ex Covellia subopposita Miq?), Bogor
(Java), Bot. Gdn., no. 9a].
Blastophaga (Ceratosolen) constricta Mayr, 1885, pp. 154, 161, 163, 169—170, Pl. XI
fig. 6 [key ®, à, descr. 9, &, ex Ficus (Cystogyne) subopposita Miq. (fistulosa Reinw.
sec. Miq.) (det. Solms), P. Mayer, no. 9a; type: 6 ].
Ceratosolen constrictus: Grandi, 1928c, pp. 184—188, figs. XXXI—XXXII [redescr. 9,
4, type specimens of B. constricta Mayr].
Material.
Series 9, &, ex Ficus fistulosa Reinw. (det. E. J. H. CORNER), Bogor (Java),
Bot. Gdn., fern garden, 1.XI.1954; coll. no. 60; &, slide 60a.
Series 9, 4, ex Ficus fistulosa Reinw. (det. E. J. H. CORNER), Bogor (Java),
Semplak, 28.X1.1954; coll. nos. 209, 211; 4, slide 21la,.9, &, 211b, ¢, 211c,
a. Dial
Series 9, 4, ex Ficus fistulosa Reinw., Tjibodas (Java), 8.11.1955; coll. no.
302.
Series 9, &, ex Ficus dimorpha King) (det. E. J. H. CORNER), Pajakumbuh
1) The wasps introduced to Hawaii, with Malayan Ficus fistulosa (cf. CORNER, 1958, p.
18), probably belong to C. hewitti Waterston.
2) — Ficus fistulosa Reinw.
O) ge I
J. T. WieBEs: Indo-Australian Ceratosolen 13
Figs. 26—38, Ceratosolen grandii, New Guinea, 26, female hind leg, 27, male mid leg, and
detail in axial aspect, 28, female antenna, axial aspect, 29, detail of female hind leg, axial
aspect, 30— 33, male, 30, mandible, 31, fore leg, 32, detail of hind leg, 33, labium and
maxillae, 34— 36, female, 34, labium and maxillae, 35, fore leg, axial aspect, and detail in
antaxial aspect, 36, mandible, 37, male antenna, 38, male. Figs. 38, X 40; 27, 31, 32, 37,
D907 26,028", 300535555) 1155129, 3273078 1.60
(Sumatra), leg. W. MEIJER, III—V.1956, no. 3473; coll. nos. 465, 467, 469; ¢,
slide 465a, 9, 4, 465b.
? Series 9, 4, ex Ficus fistulosa Reinw. var. tengerensis OK. (det. E. J. H.
CORNER), Mt. Kinabalu East (N. Borneo), 3000 ft. alt, leg. E. J. H. CORNER,
25.VII.1961, RSNB no. 967; coll. no. 626; 9, slide 626a, 4, 626b.
Description — Additional note.
The males have small eyes, near the insertion of the mandibles. Maxilla with
two lateral hairs. Head, thorax, and propodeum, fig. 2. Fore tibia with seven apical
teeth: two stout teeth at the dorsal edge, with a smaller one in between, one dor-
sally on the disk, and three more slender teeth at the ventral edge. Mid tibia
with five apical teeth. Hind tibia with three dorso-apical teeth. The aedeagus
(fig. 5) is slightly dilated, and the cerci of the tenth urite bear two large claws.
14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Female. Mandibular appendage with five or six ventral ridges. Maxilla with
one lateral hair, and a subapical one. Fore wing (2 : 1), 1.2 mm long. Submarginal,
marginal, stigmal, and postmarginal veins approximately in ratio 20 : 7 : 6 : 8. Sub-
marginal vein with three pustulus, stigmal vein with four. Hind wing (10 : 3), 0.7
mm long. Fore tibia, fig. 3; there are four apical teeth at the dorsal edge. Ventral
edge without a long spur. Mid leg long and slender. Femur two-thirds the length
of the tibia, which is as long as the tarsal segments combined. First tarsal segment
one fourth longer than the fifth, the intermediates subequal and approximately half
as long as the first. Tibia without prominent apical teeth, but with some very
stout hairs at the ventro-apical edge. Hind leg, fig. 4. Ovipositor scarcely pro-
jecting beyond the apex of the gaster.
Remark.
The specimens from Sumatra, ex F. dimorpha King, resemble the Javanese
specimens in all details. The sample from Mt. Kinabalu is provisionally referred to
C. constrictus, although the females differ slightly from the Javanese specimens:
antenna with short segments as in C. constrictus, but the sensilla rather long, as
in C. hewitti, fore tibia with two dorso-apical teeth.
Ceratosolen hewitti Waterston (fig. 1)
Ceratosolen hewitti Waterston, 1921, pp. 35—38, fig. 1 [descr. 9, 8, ex Ficus spec.,
Sarawak (Borneo), leg. J. Hewitt, VII.1907; type: 2].
Ceratosolen imbecillus Grandi, 1927a, pp. 315—317, Pl. 3 figs. 33—42, Pl. 4 figs.
43—49 [descr. 9, &, ex Ficus chartacea Wall.!), Singapore, leg. C. F. Baker]; 1928c,
p. 223 [= C. hewitti Waterston].
Material.
One 9,1 &, “from a fruiting trunk. Ficus. July 1907. J. Hewitt’, Sarawak
(Borneo), from type lot of C. hewitti Waterston; coll. BM, 1 slide.
Series 9, &, ex Ficus aff. fistulosa Reinw. (det. E. J. H. CORNER), Bau
(Kuching, Borneo), leg. E. J. H. CORNER, 25.IX.1961; coll. no. 640; 9, slide
640a, &, 640b.
Series 9, 4, ex Ficus chartacea Wall.1), Singapore, from type lot of C. m-
becillus Grandi; coll. USNM.
Series &, ex Ficus fistulosa Reinw. (det. E. J. H. CORNER), Singapore, Bot.
Gdn., 1932; coll. no. 422; &, slides 422a, b.
Series 9, 4, ex Ficus fistulosa Reinw. (det. E. J. H. CORNER), Sungei Gombah
(Selangor, Malaya), leg. E. J. H. CORNER, 4.X.1961; coll. no. 638; 9, slide 638a,
a, 638b.
Ceratosolen gravelyi Grandi
Ceratosolen gravelyi Grandi, 1916a, pp. 150, 152 [descr. 9, 4, in key]; 1916b, pp
201—207, figs. IX—XI [descr. 9, &, ex Ficus cunia Hann”), Paresnath Hill (Chota
Nagpur, India), 4300 ft. alt, leg. N. Annandale, 9 & 14.IV.1909}; Joseph, 1954, p. 409,
figs. IV, 36—41 [descr. note &, ex Ficus cunia Hann?), Dehra Dun (India), Forest
Research Inst. Estate, New Forest, leg. K. J. Joseph, 25.VI.1953].
1) Probably incorrect, see p. 97.
2) = Ficus semicordata Ham. ex Smith.
J. T. WIEBES : Indo-Australian Ceratosolen 15
Figs. 39—53, Ceratosolen boschmai, Bougainville I., 39—41, female, 39, labium and maxilla,
40, fore leg, 41, hind leg (tarsus omitted), 42, male, 43, female antenna, 44— 50, male, 44,
detail of hind leg, 45, antenna, 46, mid leg, 47, cercus of tenth urite, and paramere, 48,
aedeagus, 49, mandible, 50, labium and maxillae, 51, female mandible, 52, male fore leg, 53,
detail of female antenna in axial aspect. Figs. 42, X 40; 40, 41, 43, 44, 46, 52, 53, X 90;
39, 51, X 115; 45, 47—50, X 160
Material.
One 9,2 4, ex Ficus cunia Hann.1), Paresnath Hill (Chota Nagpur, India),
ex coll. GRANDI; coll. ML, no. 509.
Nine ¢, India, ex coll. JosEPH; coll. ML, no. 566.
1) = Ficus semicordata Ham. ex Smith.
16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Remark.
This species shows some resemblance to both C. pygmaeus and C. emarginatus,
but it appears to be more closely related to the latter.
Ceratosolen emarginatus Mayr
Ceratosolen emarginatus Mayr, 1906, pp. 153—154 [descr. 9, &, ex Ficus spec., Tonkin
(N. Vietnam), leg. P. Marchal, no. 20b}; Grandi, 1928c, pp. 189—190, figs. XX XIII—
XXXIV [redescr. 9, 4, type specimens of C. emarginatus Mayr = C. effractarius Grandi};
Joseph, 1954, p. 409 [2, &, ex Ficus roxburghii Wall), Dehra Dun (India), leg. K. M.
Vaid & K. J. Joseph, 30.VI.1953].
Ceratosolen effractarius Grandi, 1927b, pp. 169—174, figs. I-II [descr. 9, 4, ex Ficus
Roxburgi), Hanoi (Indo-China), leg. F. Silvestri, 4.1II.1925].
Material
Five 9,1 ¢, Calcutta (India); coll. USNM, incorrectly identified with Blasto-
phaga appendiculata Mayr.
Series 0 8, India, ex coll JOSEPH; coll MT. no: 56594), slides Gna,
2, 565b.
Series 9, &, ex Ficus spec., Chiangmai (N.W. Thailand), Fang, 500 m alt,
leg. T. C. MAA, 12/19.1V.1958, no. 380; coll. BMH.
Series 9, &, ex Ficus oligodon Miq.2) (det. E. J. H. CORNER), Fraser’s Hill
(Selangor, Malaya), 4000 ft. alt., leg. E. J. H. CORNER, 3.X.1961; coll. no. 609;
4, slides 609a, b, (9, 609c.
Remark.
This species is very close to C. fusctceps (Mayr). It differs, however, in having
an extra ring segment in the male antenna, and in the absence of spines from the
disk of the male hind tibia.
Ceratosolen fusciceps (Mayr)
Blastophaga spec., Mayer, 1882, p. 570 [ 9, ex Covellia glomerata Willd.?), Bogor (Java),
Bot. Gdn., no. 4a}.
Blastophaga (Ceratosolen) fusciceps Mayr, 1885, pp. 154, 161, 164, 167—168, PI. XI fig.
3 [key 9, 4, descr. 2, &, ex Ficus (Sycomorus) glomerata horti Bogor., nec Roxb. (det.
Solms), P. Mayer, no. 4a; type: 4 ].
Blastophaga (Ceratosolen) fuscipes Mayr, 1885, p. 249 [incorrect spelling}.
Ceratosolen fusciceps: Mayr, 1906, p. 153 [descr. note &, ex Ficus Covellii*), Bogor
(Java), Bot. Gdn., leg. C. Aurivillius, no. 1; also ? 9, ex Covellia glomerata Miq., and
Urostigma lucescens Miq.*), same locality, leg. C. Aurivillius, nos. 5 and 6, respectively};
Grandi, 1928a, pp. 79—80 [®, &, ex Ficus glomerata Roxb., Chambaganour (Pulneys,
India)5), 4000 ft. alt, leg. E. Gombert, 7.VI.1914]; 1928c, p. 171 [type specimens studied];
Joseph, 1953a, p. 61 [®, &, ex Ficus glomerata Roxb., Agra (India), leg. K. J. Joseph,
12.1V.1952}; 1953b, p. 137 [2, 4, ex Ficus glomerata Roxb., Agra, 28.VIII.1950, and
Trivandrum, 5.VII.1950 (India), leg. K. J. Joseph}.
GE fuscipes: Grandi, 1916a, pp. 151—152 [®, &, in key}; 1916b, pp. 194— 201,
1) = Ficus auriculata Lour.
2) The question of the host Ficus is discussed on p. 97.
3) = Ficus racemosa Linn.
4) see p. 98.
5) ? — Shembaganur, Palni Hills, South India?
J. T. WIEBES : Indo-Australian Ceratosolen 17
figs. V—VIII [descr. 9, 4, ex Ficus glomerata Roxb., Bogor (Java), Bot. Gdn., leg. E.
Jacobson, 1.1915; and Colombo (Ceylon), leg. O. Beccari, 1870]; 1923a, p. 299 [ 2, India,
leg. H. M. Lefroy].
Ceratosolen mysorensis Joseph, 1953c, pp. 277—282, figs. IV—V [descr. 9, 6, ex Ficus
mysorensis Hayne!), Trivandrum (India), Bot. Gdn., leg. K. J. Joseph, 12.V11.1950}.
?Blastophaga niveipes Girault?), 1927, p. 338 [descr. (@), ex Ficus glomeratus, Roper
River (N. Terr., Australia), leg. N. B. Tindale].
Material.
Series 9, 4, ex Ficus racemosa Linn., Bogor (Java), Bot. Gdn., VI—VIII.1954;
coll. no. 83; 4, slide 83a, 9, 4, 83b.
Series 9, 4, ex Ficus racemosa Linn., Bogor (Java), Bot. Gdn., near warung,
24.IX.1954; coll. no. 479.
Series 9, 4, ex Ficus racemosa Linn. var. elongata (King) Barrett (det. E. J.
H. CORNER), Bogor (Java), Bot. Gdn., 18.X.1954; coll. no. 236; 4, slide 236a,
2, 236b.
Series .9, &, ex Ficus racemosa Linn. (det. E. J. H. CORNER), Timor, leg. M.
E. WALSH, no. 95; coll. no. 386; 4, slide 386a, 9, 386b.
Series 9, 4, ex Ficus racemosa Linn., Sudadji (Bali, distr. Sawahan, 16 km
SE of Singaradja), leg. NJOMAN ToJA; coll. no. 59; &, slide 59a, 9, slide 59b.
Series 9, 4, ex Ficus racemosa Linn.3), Udjung Kulon (Java), leg. A. Hoo-
GERWERF, XI.1954, no. 1; coll. no. 276; 9, &, slide 276a.
Series 9, &, ex Ficus racemosa Linn.3), Udjung Kulon (Java), leg. A. Hoo-
GERWERF, XI.1954, no. 2; coll. no. 292; &, slide 292a, 9, 292b.
Series 9, &, ex Ficus racemosa Linn.3), Udjung Kulon (Java), leg. A. Hoo-
GERWERF, XI.19544); coll. no. 279; 9, 4, slide 279a.
Fifteen 9, series 4, ex Ficus racemosa Linn. var. elongata (King) Barrett5)
(det. E. J. H. CORNER), Koung, Mt. Kinabalu (N. Borneo), + 400 m alt,
17.V.1933, Sing. F. no. 27322; coll. no. 420; &, slide 420a, 9, 4, 420b.
Series 9, &, ex Ficus racemosa Linn. var. elongata (King) Barrett (det. E.
J. H. CORNER), Kota Belud (N. Borneo), leg. E. J. H. CORNER, 23.1X.1961; coll.
no. 622.
Three 9, 10 4, ex Ficus racemosa Linn. (det. E. J. H. CORNER), Sungei
Nenggiri (Kelantan, Malaya), leg. E. MCCLURE, 13.X1.1961; coll. no. 667.
Fourteen 9, 2 4, ex Ficus racemosa Linn. (det. E. J. H. CORNER), Negri
Sembilan (Malaya), 1932, Sing. F. no. 26135; coll. no. 416; 9, &, slide 416a;
do., 12 @ from a young fig; coll. no. 418.
Series 9, 7 4, ex Ficus glomerata Roxb.6), Pusa, Bihar (India), leg. D. T.
FULLAWAY, 13.II.1921; coll. HSPA.
DL) Gee jos OB
2) I saw one @ from the type series (S. Australian Museum, Adelaide), but the (as yet
unknown) male should be studied for a definitive identification. Another Australian sample
of C. fusciceps seen by me is: series 9, 4, ex Ficus glomerata Roxb. (=F. racemosa
Linn.), Greenhills, Meringa (Queensland, Australia), leg. C. E. PEMBERTON, 28.X1.1921;
coll. HSPA; coll. ML, no. 589: series 29, 4; 2, 4, slide 589a.
3) identification uncertain.
4) locality uncertain.
5) Ficus acidula King.
6) = Ficus racemosa Linn.
18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Figs. 54— 67, Ceratosolen bianchii, Fiji, 54, male fore leg, 55, female fore leg, 56, female
antenna, 57, male, 58, detail of female hind leg, 59, male hind leg, 60, female labium and
maxillae, 61—64, male, 61, detail of mid leg, 62 antenne, 63, labium and maxillae, 64,
mandible, 65, female mandible, 66, male mid leg, axial aspect, 67, apex of male abdomen.
Figs. 57, X 40; 54—56, 58, 59, 61, 66, X 90; 60, 62—65, 67, X 115
Five 9,5 &, ex Ficus glomerata Roxb. 1), Pusa, Bihar (India), leg. I. R.
DUTT, 2.111.1923; coll. HSPA; &, 2, slide mounted.
1) = Ficus racemosa Linn.
J. T. WieBEs: Indo-Australian Ceratosolen 19
One 9, 4 &, India, ex coll. JosEPH; coll. ML, no. 563; 2, slide 563a,
A IE
Two 9, 4 &, ex Ficus glomerata Roxb.1), Poona (India), leg. P. VARMA;
coll. USNM.
Ten 9,5 &, ex Ficus spec., Bangalore (India), leg. G. CoMPERE, VIII.1907;
coll. USNM.
Nine 9, ex Ficus glomerata Roxb.1), Bangalore (India), leg. D. T. FULLAWAY,
25.11.1921; coll. HSPA.
Five 9,8 &, ex Ficus glomerata Roxb.1)2), Bangalore (India), 11.1921; coll.
HSPA.
Ten 9, Peradeniya (Ceylon), leg. A. RUTHERFORD; coll. USNM.
One &, ex Ficus glomerata Roxb.1), Colombo (Ceylon), leg. STAINFORTH
GREEN; coll. OUM, 1 slide.
Ceratosolen appendiculatus (Mayr) and C. striatus Mayr
In 1885, Mayr described Blastophaga (Ceratosolen) appendiculata from the
Botanical Gardens at Bogor, Java. It was not recognized by subsequent authors,
although GRANDI, in his revision of the Agaonidae (1928c), redescribed typical
specimens from Mayr’s collections).
C. striatus Mayr was described, in 1906, as being different from C. appendicu-
latus in the shape of the head, and in the striate head and pronotum of the male.
Later, GRANDI (1917) described C. striatus notandus, the male of which is essen-
tially the same as in the nominate form, but for the non-striate head and prono-
tum, the shape of the head, and the relative length of the antennal segments. Ac-
cording to GRANDI, this new form could not be identical with C. appendiculatus,
because the female had the ovipositor three quarters the length of the gaster,
whereas in C. appendiculatus this relative length was given by Mayr as approx-
imately one third. In 1928, GRANDI referred C. striatus notandus again, with some
doubts, to the nominate form.
In my opinion, C. striatus notandus Grandi belongs indeed in the synonymy of
C. striatus Mayr, as I find characters of both (viz., the striate head and pronotum,
and the relative proportions of the antenna) in the same specimen. Moreover, after
having studied the type series of C. appendiculatus (Mayr), I am convinced of
the identity of C. striatus and C. appendiculatus. The length of the ovipositor in
the only female of C. appendiculatus before me is four-sevenths the length of the
gaster, which falls entirely within the variability of C. striatus.
Ceratosolen appendiculatus (Mayr)
Blastophaga spec. Mayer, 1882, pp. 570—571 [9, &, ex Ficus umbellata Vahl (or
umbonata Reinw.?)4), Bogor (Java), Bot. Gdn., no. 8a].
1) Ficus racemosa Linn.
2) This sample bears another label: ”F. mysorensis? COLEMAN’s comp. Bangalore”. See
the remark on Ficus mysorensis, p. 98.
3) A sample from Calcutta, in the USNM collection identified with Blastophaga
appendiculata Mayr, belongs to Ceratosolen emarginatus Mayr.
4) For a discussion of this host record, see p. 97.
20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Blastophaga (Ceratosolen) appendiculata Mayr, 1885, pp. 154, 161, 162, 164—166, PI.
XI figs. 1—2 [key 9, &, descr. 9, 6, ex Ficus (Sycomorus) umbellata horti Bogor., nec
Vahl (det. Solms)!), P. Mayer, no. 8a; type: 4 }.
Ceratosolen spec., Williams, 1928, p. 12 [biological note on the wasps from Ficus inte-
grifolia Elmer?), Philippine Is.}.
Ceratosolen appendiculatus: Grandi, 1928c, pp. 176—179, figs. XXVII—XXVIII [redescr.
2, d, type specimens of B. appendiculata Mayr].
Ceratosolen striatus Mayr, 1906, p. 153 [descr. (9), &, ex Ficus variegata BI. var.,
Bogor (Java), Bot. Gdn., leg. C. Aurivillius, no. 3}; Grandi, 1916a, p. 153 [ 4, in key};
1917, pp. 32—34, fig. XI [descr. 4, ex Ficus variegata Bl, Bogor (Java), Bot. Gdn., leg.
E. Jacobson, 1.1915}; 1928c, pp. 179—180 [type specimens of C. striatus Mayr studied;
? = C. striatus notandus Grandi}.
Ceratosolen striatus notandus Grandi, 1916a, pp. 151, 153 [@, &, in key}; 1917, pp.
34—40, figs. XII-XIII [descr. 9, &, ex Ficus variegata Bl, Bogor (Java), Bot. Gdn.,
leg. E. Jacobson, I.1915].
?Ceratosolensia ficophaga Girault?), 1915, pp. 311, 312 [descr. ®, &, ex Ficus spec.
Gordonvale (Cairns, Queensland), I, and 18.IX.1912].
Material.
One 9, 17 4, from type series of Blastophaga appendiculata Mayr; coll.
NMW.
Series 9, &, ex Ficus variegata BI., Bogor (Java), Bot. Gdn., fern garden,
2ER. 1954 :coll> n0: 222:79,, 3.7 slide M22a!
Series 9, 4, ex Ficus variegata Bl, Bogor (Java), Bot. Gdn., 24.IX.1954;
gol mor 237: Oms hels 2372.
Series 9, &, ex Ficus variegata Bl, Tjiwaringin, Bogor (Java), 22.IX.1954;
colle nos: 40, 125519. 8, slide 40a:
Series 9, &, ex Ficus variegata BI. (det. E. J. H. CORNER), Tjibodas (Java),
30.X1.1954 colli 102230: 907, 33 slide 230a.
Fragments 9, 4, ex Ficus variegata BI. var. garciae (Elmer) Corner (det. E. J.
H. CORNER), Mt. Iraya (Batan I., Philippine Is.), Bur. Sci. no. 80191; coll.
no. 430; ®, &, slide 430a.
Series 2, &, ex Ficus integrifolia Elmer?), Los Baños (Luzon, Philippine Is.),
leg. EF. Xe WALLIAMS. 17.V.1921 coll ASPAS col SMIS n035952 32.27 3 slide
593a.
Six 4, several immature 9, ex Ficus variegata BI. var. sycomoroides (Miq.)
Corner (det. E. J. H. CORNER), Mt. Maquiling (Luzon, Philippine Is.), Coll.
Agric., Laguna, no. 592; coll. no. 409; 4, slides 409a, b, immature 9, 409c.
Four 9, 4 &, ex Ficus variegata Bl, Mt. Maquiling (Luzon, Philippine Is.),
“on the Calamba-trail”, leg. F. X. WILLIAMS, 30.1.1922; coll. HSPA.
Series 9, &, ex Ficus variegata Bl, Mt. Maquiling (Luzon, Philippine Is.),
1) For a discussion of this host record see p. 97.
2) = Ficus variegata Bl. var. sycomoroides (Miq.) Corner.
3) I saw 1 @ from the type series (Queensland Museum, Brisbane), but the male should
be studied for a definitive identification. Other Australian samples of C. appendiculatus
seen by me are:
Series @, &, ex Ficus ehretioides F. v. Muell. (= Ficus variegata BI.), Hambledon
(Queensland), leg. C. E. PEMBERTON, 6.XII.1921; coll. HSPA.
Series Q, 4, ex Ficus ebretioides F. v. Muell. (= Ficus variegata BI.), Merawa
(Queensland), leg. C. E. PEMBERTON, 18.X1.1921; coll. HSPA.
J. T. WIEBES : Indo-Australian Ceratosolen 21
leg. F. X. WILLIAMS, 11 & 14.XII.1921; coll. HSPA; coll. ML, no. 592: 5 9,
DOr
Immature 9, &, ex Ficus viridicarpa Corner!) (det. E. J. H. CORNER), Ke-
manan (Trengganu, Malaya), 23.X1.1935, Sing. F. no. 30553; coll. no. 478;
immature 9, &, slide 478a.
Immature 9, &, fragments, ex Ficus viridicarpa Corner!) (det. E. J. H. CORNER,
type no.), Jelebu (Negri Sembilan, Malaya), 29.X.1932, Sing. F. no. 26032; coll.
no. 432; immature 9, ¢, slide 432a, ¢, fragments, 432b.
Ceratosolen grandii sp. n. (figs. 26—38)
Material.
Series ¢, a few immature (9, ex Ficus semivestita Corner (det. E. J. H.
CorNER, type no.), Morobe district (Terr. New Guinea), 3000 ft. alt., leg. E. J.
H. CORNER, 10.X.1955, NGF no. 7358; coll. no. 452; & (holotype), slide 452a,
2 (allotype), 452b.
Description.
Male. Head (fig. 38) nearly twice as long as wide anteriorly, and longer than
its maximum width [6 : 5}, with many small triangular bristles. The lateral lobes
of the epistomal margin are much longer than the median one, and bear small
hairs. Eyes present. Antennal grooves half closed. Antenna (fig. 37) four-
segmented. Pedicel (2:1) two-thirds the length of the scape (2:1), third seg-
ment (2:1) somewhat longer than the pedicel, and approximately half as long as
the apical segment (4:1). Mandible, fig. 30. Labium and maxillae, fig. 33; the
maxilla bears two subbasal hairs.
Thorax, fig. 38. Pronotum only little longer than wide anteriorly, its maximum
width one and a half times the length. Mesonotum wide, the length three-fifths of
the width. Metanotum anteriorly nearly as wide as the mesonotum, narrower be-
hind. Propodeum narrow and rather long, indistinctly separated from the meta-
notum. Tibia of the fore leg (fig. 31) half as long as the femur, tarsus nearly as
long as the tibia. Tibia with two dorsal and three ventro-apical teeth; a third dor-
sal tooth is situated on the disk. Tarsal segments incompletely separated, the first
with three conical ventral spines; the second, subequal in length to the first (meas-
ured dorsally), without spines. Mid leg, fig. 27. Tibia with one ventro-apical
tooth, about as long as the femur, and but little shorter than the tarsus. Dorsal
surface of the tibia provided with diverging rows of conical spines; one spine at
the ventral margin. First tarsal segment nearly as long as the three following
combined, with a pair of ventral spines. Fifth segment shorter than the first
[2 : 3}, but about twice as long as the intermediate segments. Tibia of the hind leg
(fig. 32) with a bidentate ventro-apical tooth; the disk with hairs that are situated
on small protuberances. First tarsal segment as long as the three following combin-
ed, with seven ventral conical spines. Second to fourth segments with three apical
spines each. Fifth segment shorter than the first [3 : 5}, without spines.
n) SEE fo Sie
22 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Figs. 68—79, Ceratosolen armipes, New Guinea, 68—71, female, 68, detail of fore leg, 69,
antenna, 70, detail of mid tibia, 71, hind leg, 72—78, male, 72, antenna, 73, male, 74, mid
leg, 75, cercus of tenth urite, 76, detail of fore leg, 77, mandible and hypostomal margin,
78, detail of hind leg, 79, female mandible. Figs. 73, X 40; 68—71, 74, 76, 78, 79, X 90;
Ue Vs DR USE UD AK 275
Gaster. Genitalia without cerci.
Length, 1.6 mm. Colour yellowish brown.
Female. The description is made from immature specimens taken from the gall-
flowers. Head slightly longer than wide across the compound eyes. Longitudinal
diameter of the eye one and a half times as long as the cheek. Antenna (fig. 28)
with nine free segments. Scape large, twice as long as the pedicel. Flagellum short,
with rather wide segments. Funicular segments with one row of sensilla each; the
club, consisting of the apical three segments, with one complete and one in-
J. T. WIEBES : Indo-Australian Ceratosolen 23
complete row. Mandible, fig. 36: its appendage with four transverse ridges.
Labium and maxillae (fig. 34) with one apical hair each.
Thorax very much like that of C. appendiculatus, but scutum and scutellum
with more hairs: eight and twenty, respectively. Close to the propodeal spiracle
there are only a few hairs. Wings still folded; not studied. Femur of the fore leg
(fig. 35) nearly as long as tibia and tarsus together. Tibia with four apical teeth
visible in antaxial aspect. First tarsal segment not quite as long as the three fol-
lowing combined, second to fourth segments subequal, the fifth as long as third
and fourth segments combined. Tarsal segments with stout ventral hairs: ten on
the first, others with two apicals only, except for the fifth, which has three ven-
trals. Tibia of the mid leg nearly as long as femur and trochanter combined,
without apical spines, but with many spines on the disk. Tarsal segments in ratio
8:5 :4:3:8 (with the claws). Hind leg, figs. 26, 29. Coxa and femur subequal,
length of the tibia two-thirds of that of the femur. Tibia with two apical bidentate
teeth. Tarsal segments in ratio 14:5 : 6 : 4 : 6; all segments with rows of ventral
spines (fig. 29).
Gaster. Protruding part of the ovipositor approximately half as long as the
gaster.
Length, 1.6 mm.
Remark.
This species appears to be close to C. appendiculatus, judging from the structure
of the female antenna, and of the male genitalia. On the other hand there are
several structures in the male suggesting a close affinity to C. fusciceps: antenna
consisting of four segments, mouthparts not atrophied.
I dedicate this species to Professor GUIDO GRANDI (Bologna), the well-known
authority on fig wasps.
Ceratosolen boschmai sp. n. (figs. 39—53)
Material.
Series 9, &, ex Ficus salomonensis Rech. (det. E. J. H. CORNER), Arawa,
Kieta (Bougainville I.), leg. E. J. H. CORNER, X.1960; coll. no. 532; & (holo-
type), slide 532a, 9 (allotype), 532b, 4, 9 (paratypes), 532c, d.
Description.
Male. Head (fig. 42) approximately twice as long as wide anteriorly, maximum
width two-thirds of the length. Dorsal surface with many small hairs. Lateral lobes
of the epistomal margin with one apical hair. Eyes wanting. Antennal grooves half
closed. Antenna (fig. 45) five-segmented; the scape (3 : 2) about twice as long
as the pedicel (5:4), which is not quite twice as long as the next segment
(5 :3). This segment is, in some specimens, half split into two parts, in others
it is entire, or shows only a small notch at the antaxial margin. Fourth segment
(7 : 4) three to four times as long as the third, the apical segment (2:1) thrice
as long as the third. Mandible, fig. 49. Labium and maxillae (fig. 50): there are
three (one basal and two lateral) hairs on each maxilla.
24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Thorax, fig. 42. The pronotum is rather narrow, and does not quite conceal the
propleurae. Posteriorly it is about twice as wide as it is anteriorly; its length is
equal to the sum of the anterior and posterior widths. Mesosternum about as long
as wide, with rounded lateral edges. Metanotum consisting of two ear-like plates,
not quite separated in the mid line, and almost free from the long propodeum.
The lateral edges of the propodeum are folded ventrally, leaving a very narrow
dorsal surface; the lateral surfaces are almost wholly occupied by the large peri-
tremata. The fore leg (fig. 52) shows a very long femur, much longer than tibia
and tarsus combined [3 : 2}. Tibia (without the apical teeth) more than twice as
long as the second tarsal segment, which is slightly longer than the first [6 : 5].
Tibial armature consisting of four dorsal and two ventro-apical teeth. Tarsus
bimerous, the segments incompletely separated. Mid leg, fig. 46. Tibia as long as
femur and trochanter combined, and but little longer than the pentamerous tarsus.
Tarsal segments in ratio 7:3 :3 :3 :5. Femur of the hind leg (fig. 44) large,
tibia not much more than half as long, with three dorso-apical teeth. Tarsus
pentamerous, segments in ratio 9:3:3:2:6.
Tergites of the gaster distinctly sclerotized. Genitalia (fig. 48) with cerci (four
claws) and parameres (fig. 47).
Length, 1.5—1.9 mm. Colour yellowish brown, head and fore legs darker, the
propodeal peritremata strikingly black-brown.
Female. Head approximately as long as wide across the eyes. Longitudinal diam-
eter of the eye short, about four-fifths the length of the cheek. Lateral lobes of
the epistomal margin rather angular. Antenna (fig. 43): the scape two and a half
times as long as the pedicel, which has many minute spines on the axial surface
(fig. 53). Fifth to eleventh segments with one row of oblong sensilla; the fifth
with five sensilla, the sixth with six, the seventh and eighth with eight, the ninth
with nine, the tenth with ten, and the eleventh with four oblong and two circular
sensilla. Ninth to eleventh segments combined into a loose club. Mandible (fig.
51) with four ventral ridges, its appendage with five. Labium and maxilla, fig. 39.
The bacilliform process is about half as long as the maxilla.
Thorax very finely striate, with long hairs. Scutum not quite as long as its
maximum width [3 :4}, with fifteen hairs. Scutellum about as long as wide,
with seven hairs on each side. Metanotum not much wider than scutellum and
axillae combined, propodeum distinctly wider. Propodeal peritremata large; tufts
of about ten hairs occur beneath the spiracles. Fore wing (9 : 4), 1.6 mm long;
nearly the whole surface setose. Submarginal, marginal, stigmal, and postmarginal
veins in ratio 22 : 6 : 7 : 11; stigmal vein with three or four pustules, submarginal
vein with three. Hind wing (5 : 2), 1.0 mm long. Femur of the fore leg (fig. 40)
about as long as coxa and trochanter combined, the tibia half as long, with five
apical teeth — the first of which is blunt — arranged in a dorsal comb. Tarsal
segments with expanded plantar edges, the first segment is distinctly longer than
the fifth [7:5}, and more than twice as long as the subequal intermediates.
Tibia of the mid leg about as long as trochanter and femur combined. Tarsus
pentamerous, the segments in ratio 17:7:8:7:14. Femur of the hind leg
(fig. 41) with a very distinct ventral groove. Tibia with one apical axial tooth,
J. T. WieBEs: 1ndo-Australian Ceratosolen 25
Figs. 80—83, Cera:osolen abnormis, West New Guinea, male, 80, detail of hind leg, 81,
apex of abdomen, 82, mandible, 83, mid leg, axial aspect, 84— 91, C. sordidus, Solomon I,
84, detail of female hind leg, 85—88, male, 85, mandible, 86, detail of hind leg, 87, male,
88, mid leg, 89, detail of female fore leg, 90, male antenna, 91, male fore leg. Figs. 87,
X 40; 81, 91, X 90; 80, 82—86, 88—90, X 115
26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
and a bidentate antaxial tooth. Tarsal segments in ratio 20:8:5:6:11.
Gaster. Ovipositor distinctly projecting.
Length, 1.7—2.1 mm. Colour black-brown, distal parts of the extremities
lighter.
Remark.
C. boschmai is easily distinguished from its congeners by the peculiar propo-
deum of the male. It is a pleasure to dedicate this species to Dr. H. BOSCHMA,
Professor of Systematic Zoology at Leiden University.
Ceratosolen bianchii sp. n. (figs. 54—67)
Material.
Series some MEENDEN (no 27), Fiji 1. MES MC ES PEMBERMON 1920,
coll. HSPA, slide mounted: & (holotype), ® (allotype), 9, & (paratypes);
coll MESN0059825, OR 2 7a
DESC DETO DL
Male. Head (fig. 57): the width two-thirds of the length. Median lobe of
epistomal margin broad. Eyes present. Antennal grooves half closed. Antenna
(fig. 62) five-segmented; the scape not quite twice as long as its maximum width,
the pedicel as long as the width of the scape, and half as wide. Third segment
(1:2) annuliform, fourth and fifth segments subequal in length, the latter
(2:1) is narrower than the former. Labium and maxillae, fig. 63; labium with
one ventral hair, maxilla with one lateral hair. The maxillae are rather long; and
they fold easily in the slides. Mandible, fig. 64.
Thorax, fig. 57. Pronotum approximately twice as long as wide anteriorly, wider
behind. Mesonotum large; metanotum narrower, incompletely separated from the
long propodeum. Propodeal peritremata large. Femur of the fore leg (fig. 54)
more than twice as long as the tibia. Apical dentation of the tibia consisting of
four dorsal and two ventro-apical teeth. Tarsus bimerous, though the two seg-
ments are not completely separated. Segments in ratio 5 : 7. Mid leg, figs. 61, 66.
Femur longer than coxa and trochanter combined, but shorter than the tibia
{8:11}. Tibia with a few stout hairs and spines, especially on the antaxial sur-
face, and one ventro-apical tooth. Tarsus pentamerous, segments in ratio
9:3:4:3:12. Coxa of the hind leg (fig. 59) semiglobular, trochanter small;
the femur expanded dorsally. Tibia not quite as long as the femur, with four
antaxial ventro-apical teeth. Tarsus pentamerous, the segments in ratio 24:9:8:
SERIE
Gaster. Genitalia (fig. 67) without cerci or parameres; apex of the aedeagus
scarcely dilated.
Length, 2.0—2.1 mm. Colour uniformly dark yellow-brown.
Female. Head approximately as long as wide across the eyes. Eyes small, the
longitudinal diameter distinctly shorter than the cheek [7 :9}. Face with rather
1) As the material has been desiccated, some characters are not easily to be observed, and
some must remain undescribed.
J. T. WIEBES : Indo-Australian Ceratosolen 27
long pubescence. Antenna, fig. 561). Scape two and a half times as long as the
pedicel. Fifth segment with five oblong subapical and two circular sensilla. Sixth
segment twice as long as the fifth, seventh and eighth segments subequal, shorter
than the sixth. Apical three (or two?) segments shaped so as to form a club, ninth
and tenth segments subequal, the eleventh shorter. Sixth to eleventh segments with
irregular rows of oblong sensilla. Mandible (fig. 65) with twelve ventral ridges,
its appendage with ten. Labium and maxillae, fig. 60. Labium with one apical
hair, maxilla with a subapical hair and a bacilliform process. The latter has not
quite one third the length of the maxilla, and bears a long apical hair.
Thorax pubescent, relative proportions much as in C. boschmai. Propodeum with
five hairs below the large peritremata. Fore wing, 2.0 mm long, more than twice
as long as wide. Submarginal, marginal, stigmal, and postmarginal veins in ratio
15 :6:4:6. Stigmal vein with four pustules, submarginal vein with three. Post-
marginal vein rather robust and dark. Hind wing (6:1), 1.3 mm long. Fore leg
(fig. 55): femur and trochanter together as long as tibia and tarsus combined, and
not quite twice as long as the coxa. Apical dentation of the tibia consisting of
five teeth, the ventral one of which is blunt; the dorsal one is the longest. Tarsus
five-segmented; first and fifth segments subequal in length, fourth segment approx-
imately half as long, second and third segments shorter, subequal. Coxa of the mid
leg semiglobular; the trochanter small, the femur thrice as long; the tibia a little
longer than the femur, with one ventro-apical spur. Tarsal segments in ratio
42:20:21 :16:25. Hind leg, fig. 58. Femur with a distinct ventral groove.
Antaxial apical tooth of the tibia tridentate (the median tooth the longest); the
axial apical tooth long and curved. Tarsal segments in ratio 15 :7:5:4:7, rather
pubescent.
Gaster. Ovipositor slightly projecting beyond the apex of the gaster.
Length, 2.3 mm. Colour olive-brown, thorax and apical part of the gaster
darker.
Remark.
This species appears to be close to C. boschmaz, but it differs in the relative
proportions of the male thorax, the absence of cerci and parameres from the male
genitalia, the female antenna, the dentation of the fore and hind tibiae, etc.
I have great pleasure in naming the new species after F. A. BIANCHI, Principal
Entomologist, Experiment Station, HSPA, Honolulu, who gave much help by
making F. X. WırLıams’ collection of fig wasps available for examination.
Ceratosolen armipes sp. n. (figs. 68—79)
Material.
Series 9, 4, ex Ficus itoana Diels (det. E. J. H. CORNER), Wau (Terr. New
Guinea), 3000 ft. alt, leg. E. J. H. CORNER, 13.IX.1960; coll. no. 546; &
(holotype), slide 546a, 9 (allotype), 546d, 9, & (paratypes), 546b, c, e.
reisten pt v0):
Male. Head (fig. 73) nearly rectangular, longer than wide [5 : 4]. Dorsal
28
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Figs. 92—103, Ceratosolen abnormis, West New Guinea (94, 103: specimens from East
New Guinea), 92—96, female, 92, detail of fore leg, 93, fore tibia, axial aspect, 94, head,
95, antenna, 96, detail of hind leg, 97, male antenna, 98, male, 99, female mandible, 100,
female labium and maxillae, 101—103, male, 101, detail of fore leg, axial aspect, 102,
apex of fore tibia, 103, labium and maxillae, lateral aspect, 104—105, C. bakeri, Luzon,
male, 104, labium and maxillae, 105, apex of abdomen. Figs. 94, 98, X 40; 95, X 90;
92, 93, 96, 97, 99— 103, 105, X 115; 104, X 160
J. T. WIEBES : Indo-Australian Ceratosolen 29
surface anteriorly with small hairs. Epistomal margin broadly tridentate, the
lateral lobes with one subapical hair, and one hair at mid length. Antennal grooves
half closed. Antenna (fig. 72) four-segmented. Pedicel (2:1) two-thirds the
length of the scape (2:1), and little shorter than the third segment (2:1).
Apical segment (14 : 5) nearly one and a half times as long as the third segment.
Mandible, fig. 77. Labium and maxillae atrophied, the hypostomal margin with
two long hairs.
Thorax, fig. 73. Pronotum approximately twice as long as wide anteriorly, the
posterior width two-thirds of the length. Mesonotum sub-oval, wider than long
[5 : 4]. Metanotum narrower than the mesonotum, about twice as wide as long.
Propodeum incompletely separated from the metanotum, longer than wide [13 : 9},
with the large spiracular peritremata protruding laterally. Fore leg, fig. 76. Femur
twice as long as the tibia. Tibia with three dorsal and three ventro-apical teeth.
Tarsus bimerous, segments in ratio 7:8. Tibia of the mid leg (fig. 74) longer
than the femur [5 : 4], the ventro-apical edges produced; the dorsal edge with
many stout spines, which also occur on the axial and antaxial disks, and at the
ventral apex. Tarsus pentamerous, segments in ratio 14:5 :5:5:16. Tibia of
the hind leg (fig. 78) shorter than the large femur {8 : 11], with three ventro-
apical teeth. Dorsal edge produced at mid length. Tarsus pentamerous, segments
in ratio 21:7:5:5:19, with ventral protuberances and few hairs.
Gaster. Tergites weakly sclerotized. Genitalia with very small, hyaline cerci
(fig. 75), which seem to bear two claws.
Length, 2.2—2.3 mm. Colour dark brown.
Female. Head approximately as long as wide across the eyes. Longitudinal diam-
eter of the eye not quite as long as the cheek. Pubescence scattered, denser
towards the inner rims of the eyes. Antenna (fig. 69) rather short, the scape
large, nearly thrice as long as the pedicel. Axial surface of the pedicel with about
one hundred and fifty small spines. Fifth segment with six short sensilla. The
other segments bear two to three irregular rows of small sensilla, and they are
heavily pubescent. Sixth to ninth segments subequal, tenth and eleventh segments
united into a club. Mandible (fig. 79), and its appendage, with eight ventral
ridges. The labium bears two long apical hairs.
Thorax. Pronotum wide. Scutum as long as wide at mid length, wider anteriorly,
and tapering behind. Width of the scutellum four-fifths of the length; scutellum
with about twenty hairs. Metanotum with a few lateral hairs. Propodeum with ten
hairs next to the spiracular peritremata, and four larger hairs beneath. Fore wing
(2:1), 2.1 mm long. Submarginal, marginal, stigmal, and postmarginal veins in
ratio 9:3 :2:6. Submarginal vein with two pustules, stigmal vein with three.
Hind wing (7 : 2), 1.2 mm long. Wings pubescent. Fore leg, fig. 68. Femur more
than one and a half times as long as the coxa, and nearly twice as long as the
tibia. Tibia apically with a dorsal comb of five teeth, the first of which is blunt.
Tarsus pentamerous, rather pubescent, segments in ratio 5:2:2:1:3. Tibia of
the mid leg nearly as long as trochanter and femur combined, with two apical
spurs (fig. 70). Tarsal segments in ratio 16:7:6:4:8. Hind leg, fig. 71: the
femur about one and a half times as long as the coxa, and nearly twice as long
30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
as the tibia. Ventral edge of the femur with a distinct groove. Apex of tibia with
a shovel-shaped axial tooth and a three-dentate antaxial tooth. Tarsus pubescent,
segments in ratio 16:7:6:5 :7.
Gaster. Projecting part of the ovipositor approximately one third the length of
the gaster.
Length, 2.0—2.3 mm. Colour dark brown, with lighter extremities.
Remark.
C. armipes is remarkable in having, among other differential characters, the
armature of the male mid leg rather heavy, the propodeum of a peculiar shape, and
the apical two segments of the female antenna united into a club. Apparently, it
is related to C. boschmai, C. bianchit, and C. sordidus, and to some extent also to
C. abnormis.
Ceratosolen sordidus sp. n. (figs. 84—91)
Material.
Series 4, 1 immature 9, ex Ficus cynaroides Corner (det. E. J. H. CORNER),
Bougainville I. (Solomon Is.), leg. J. K. L. WATERHOUSE, 1933, no. 845;
coll. no. 552; 4 (holotype), slide 552b, & (paratype), 552a, 9, 4 (allotype
and paratype, in copula), 552c.
Dieisicn ip eto:
Male. Head (fig. 87): width at mid length three quarters of the length, slightly
narrower in front and behind. Lateral lobes of the epistomal margin wide. Eyes
absent. Antennal grooves half closed. Antenna (fig. 90) five-segmented; the scape
(9 : 5) one and a half times as long as the pedicel (2 : 1), third segment annuli-
form. Fourth segment (15 : 8) shorter than the apical segment (19:8), which
is approximately as long as the scape. Mandible, fig. 85. Labium and maxillae
present, but not clearly visible in the badly preserved material.
Thorax, fig. 87. Pronotum nearly twice as long as wide anteriorly, and not
much longer than wide posteriorly. Mesonotum transverse, incompletely separated
from the metanotum, and about as wide. Metanotum and propodeum narrowly
fused; propodeum more than twice as long as wide, with large spiracular peritre-
mata. Fore leg, fig. 91. Femur more than twice as long as the tibia, which bears
two apical teeth at the ventral edge, and four at the dorsal edge. Tarsus bimerous,
segments subequal. First segment with small ventral spines. Femur and tibia of
the mid leg (fig. 88) subequal in length; the tibia with a few dorsal spines, the
ventral edge produced. Tarsus pentamerous, with stout ventral spines on the first
four segments; the segments in ratio 10 : 3 : 4 : 3 : 8. Tibia of the hind leg (fig.86)
with four apical teeth, and with small spines along the dorsal edge. First four
tarsal segments with ventral spines and small protuberances; the segments in ratio
IS 021
Gaster. The genitalia seem to bear no cerci, but the state of preservation does
not permit of a close examination.
Length, 1.0—1.1 mm.
J. T. WieBES: Indo-Australian Ceratosolen 31
Figs. 106—116, Ceratosolen gracilis, Luzon, 106, male hind leg, 107, apex of male mid
tibia, 108, detail of female fore leg, 109, male fore leg, 110, female mandible, 111—114,
male, 111, labium and maxillae, 112, antenna, 113, male, 114, apex of abdomen, 115,
detail of female hind leg, 116, male mandible. Figs. 113, X 50; 106, X 80; 109, X 115;
107, 110, 112, 115, X 155; 108, 114, 116, X 215; 111, X 365
Female. The immature female specimen is badly preserved. Some characteristics
are: the antennal club consists of the apical three segments, which, however, are
not completely fused; the fore tibia (fig. 89) and the fore and hind tarsi are
heavily armed with spines; the hind tibia (fig. 84) bears an apical armature,
which is shaped as in the other species related to C. armipes.
Remark.
Although the specimens are badly preserved, and the new species could not
be described in full, C. sordidus is easily recognized by the shape of the male
head and thorax, and by the armature of male and female legs.
32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Ceratosolen bakeri Grandi (figs. 104 and 105)
Ceratosolen bakeri Grandi, 1927a, pp. 312—314, Pl. 2 figs. 18—25, Pl. 3 figs. 26—32
{descr. 9, &, ex Ficus pseudopalma Blanco, Los Baños (Luzon, Philippine Is.), leg. C. F.
Baker}; Williams, 1928, pp. 12—13 [biological note}.
Material.
Series 9, 4, ex Ficus pseudopalma Blanco, Los Baños (Luzon, Philippine Is.),
leg. C. F. BAKER; coll. USNM.
Series 9, &, ex Ficus pseudopalma Blanco, Manila (Luzon, Philippine Is.),
leg. WESTER, 30.IV.1919; coll. HSPA; coll. ML, no. 591: 2 9,2 6.
Fragments of 2 9, ex Ficus pseudopalma Blanco (det. E. J. H. CORNER),
Phil. Nat. Herb. no. 16894; coll. ML, slide no. 385a.
Description — Additional note.
Male. Labium and maxillae (fig. 104) with one long hair each. Genitalia (fig.
105) without cerci.
Ceratosolen abnormis sp. n. (figs. 80—83; 92—103)
Material.
Two immature 9, 6 &, ex Ficus dammaropsis Diels (det. E. J. H. CORNER),
Begowri River (N.W. New Guinea), leg. GJELLERUP, no. 207; coll. no. 408; &
(holotype), slide 408a, 9 (allotype), 408d, & (paratypes), 408b, c.
One immature 9, ex Ficus dammaropsis Diels (det. E. J. H. CORNER), Albatros
Bivak (West New Guinea), leg. W. M. DOCTERS VAN LEEUWEN, no. 9162;
coll. no. 394.
Three 2, series &, ex Ficus dammaropsis Diels var. obtusa Corner, Aiyura
(Terr. New Guinea), Highland Experiment Station, leg. J. H. ARDLEy, 5.IX.
WSs COL no S753 2, @ (patatypes)), slides 5752, bye
Design ipo:
Male. Head (fig. 98) one third longer than its maximum width and a little
over twice as long as wide anteriorly. Eyes absent. Epistomal margin with two
large, wide lateral lobes; the median lobe is smaller. Lateral lobes with many small
hairs. Antennal grooves half closed. Antenna (fig. 97) five-segmented; scape
(15 :8) one and a half times as long as the pedicel (5 : 3); third segment an-
nuliform. Fourth segment (4:3) shorter than the pedicel, and about one third
the length of the apical segment (11 :3). Mandible, fig. 82. Labium and maxil-
lae (fig. 103); labium with one (coll. no. 575) or three (coll. no. 408) hairs.
Thorax, fig. 98. Pronotum little longer than wide posteriorly. Anterior width
nearly three quarters of the length. Mesonotum very wide, distinctly more than
twice as wide as long. Metanotum narrower, about as wide as the pronotum
posteriorly, incompletely separated from the propodeum. Propodeum approximate-
ly as long as wide, with large peritremata. Fore leg, figs. 101, 102. Femur nearly
twice as long as the tibia. Tibia with four dorso-apical teeth, and two ventral
J. T. WIEBES : Indo-Australian Ceratosolen 33
teeth. Tarsal segments fused, with a small ventral notch reminding one of the
usual bimerous condition. Trochanter of the mid leg (fig. 83) half as long as
the coxa, the femur not quite as long as the tibia. Tarsus pentamerous, segments
in ratio 32 : 12 : 11 : 18 : 50. Coxa and tibia of the hind leg (fig. 80) subequal in
length, the femur is considerably longer. Tibia with three apical teeth at the
ventral edge, the relative proportions of which vary in the various samples. Tarsus
with stout ventral hairs; first segment approximately thrice as long as the second,
second to fourth segments subequal, the fifth segment, including the claws, two
and a half times as long as the first.
Gaster. Tenth urite, and aedeagus: fig. 81. The genitalia are different from the
normal type in having very large parameres. The apodemae aedeagales are rather
short and robust.
Length, 2.2 mm. Colour dark brown.
Female. The description was originally made from immature specimens (coll.
no. 408), but it was later completed from the mature specimens of sample no. 575.
Head (fig. 94) approximately as long as wide across the eyes. Longitudinal
diameter of the eye slightly shorter than the cheek. Antenna (fig. 95) rather
short. Scape large, thrice as long as the pedicel. Third segment rather robust,
fourth and fifth segments long and slender, the fifth with circular sensillar pits in
the apical half. Sixth segment without sensilla, seventh segment with one oblong
sensillum. Eighth and ninth segments wide, with two incomplete, irregular rows
of sensilla; apical two segments united into a club, with irregular rows of sensilla.
Mandible (fig. 99), and its appendage, with six ventral ridges. Labium with two
hairs: maxillae in most specimens without hairs, in one specimen (575a) there is
one lateral hair on one side only (fig. 100).
Thorax. Pronotum large, approximately twice as wide as long. Scutum not
much shorter than wide anteriorly, glabrous. Scutellum as long as wide posteriorly,
narrower in front; with about fifteen to twenty hairs. Metanotum short, lateral
parts longer, with fifteen hairs. Propodeum large, nearly half as long as wide;
spiracles concealed from dorsal view by antero-lateral expansions. Many long hairs
occur next to and beneath the peritremata. Fore wing (5 : 2), 2.9 mm long. Sur-
face pubescent, except for some glabrous parts near the base of the wing; with
indistinct darker striae radiating from base and stigma. Submarginal, marginal,
stigmal, and postmarginal veins in ratio 24 : 6 : 6 : 13. Postmarginal vein distinctly
enlarged apically. Submarginal and stigmal veins with three pustules each. Hind
wing (4:1), 1.65 mm long. Fore leg (fig. 92): femur not quite as long as tibia
and tarsus combined. The tibia bears an apical row of eight to ten teeth, the
dorsal one of which is particularly long. As seen on the axial side (fig. 93), the
tibia appears to be a very slender segment, dorso-antaxially broadened into the
wing-shaped shield bearing the apical teeth. Tarsal segments in ratio 14:5 :6:9:
11. Tibia of the mid leg not much longer than the femur, and as long as the
tarsus. Tarsal segments in ratio 10 : 6 :5 :5 :8 (including the claws). Hind leg,
fig. 96. The femur has a distinct ventral groove. Apically, the tibia bears a quad-
tridentate appendage at the antaxial edge, and a spade-shaped tooth at the axial
edge. Tarsal segments in ratio 14:5 :4:3 : 6 (including the claws).
34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Figs. 117—129, Ceratosolen hooglandi, New Guinea, 117, female labium and maxillae,
118—122, male, 118, hind leg, 119, labium and maxillae, 120, fore leg, axial aspect, 121,
antenna, 122, mandible, 123—125, female, 123, stigmal vein of fore wing, detail, 124,
antenna, 125, mandible, 126, apex of male abdomen, 127, male, 128, female fore tibia,
129, detail of female hind leg. Figs. 127, SC GOR ile, WO, DE ODP IDA, SK 1953 A7, 125%
1250128 129) 0185 012101225126, 00255 211000315
J. T. WieBEs: Indo-Australian Ceratosolen 35
Gaster. The projecting part of the ovipositor three quarters the length of the
gaster; the sheaths robust.
Length, 2.35 mm. Colour black-brown; wings infuscated.
Remark.
The female of C. abnormis is abnormal in various aspects (e.g., the antenna,
dentation of fore and hind tibiae), but the species shows some resemblance to
C. armipes and relatives.
There is some variation between the specimens from West and East New
Guinea, to which more attention should be given when more and better material
becomes available.
Ceratosolen crassitarsus (Mayr)
Blastophaga (Ceratosolen) crassitarsus Mayr, 1885, pp. 154, 161, 163, 171—172, PI. XI
fig. 7 [key 2, &, descr. 9, &, ex Ficus (Cystogyne) Ribes Reinw. (det. Colms), Sukawana
(Tangkuban Prahu, near Bandung, Java), leg. H. Solms-Laubach; type: 4 ].
Ceratosolen crassitarsus: Grandi, 1916a, pp. 150, 152 [®, &, in key, Java]; 1917, pp.
40—46, figs. XIV—XV [descr. 2, 4, ex Ficus ribes Reinw., Tjibodas (Java), Bot. Gdn.,
leg. J. Boldingh, 11.1915}; 1928c, p. 172 [type specimens studied].
Material.
Six 9, ex Ficus ribes Reinw., Tjibodas (Java), leg. J. H. DE GUNST, 5.X1.1954;
coll. no. 94.
Series 9, 4, ex Ficus ribes Reinw., Tjibodas (Java), 20 & 23.XII.1954; coll.
no. 145; ¢, slide 145a.
Remark.
For the records of C. ? crassitarsus: Grandi (1923a, p. 299; 1927b, p. 174),
SEE) Pp. 83.
Ceratosolen gracilis sp. n. (figs. 106—116)
Material.
Series ¢, ex Ficus ribes Reinw. var. cuneata (Miq.) Corner (det. E. J. H.
CoRNER), Mt. Pinatubo (Prov. Pampanga, Luzon, Philippine Is.), ELMER no.
21980; coll. no. 401; 4 (holotype), slide 401a, 4 (paratypes), 401b.
Series 9, ¢, ex Ficus ? merrilli1), Mt. Maquiling (Luzon, Philippine Is.), leg.
F. X. WILLIAMS, 11.V.1921, no. 202; coll. HSPA, 9 (allotype and paratypes),
and & (paratypes) slide mounted; coll. ML, no. 681: 10:9, 10 4.
Series 9, &, ex Ficus merrillit), Mt. Maquiling (Luzon, Philippine Is.), leg.
F. X. WILLIAMS, 11.V.1921, no. 303; coll. HSPA, @ (paratype) slide mounted.
Series 9, 3, ex Ficus ribes2), Mt. Maquiling (Luzon, Philippine Is.), leg. F.
X. WILLIAMS, 10.V.1921, no. 404; coll. HSPA.
Seven 9, 6 6, ex Ficus ribes2), Mt. Maquiling (Luzon, Philippine Is.), leg.
F. X. WILLIAMS, 26.XI.1921; coll. HSPA.
1) Ficus merrilli Elmer = F. ribes Reinw. var. cuneata (Miq.) Corner.
2) True Ficus ribes Reinw. does not occur in the Philippines.
36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Dye sie riipiti om’.
Male. Head (fig. 113) more than twice as long as wide anteriorly, its maximum
width nearly two-thirds of the length. Lateral lobes of the epistomal margin rather
sharp. Pubescence sparse and short. Eyes distinct. Antennal grooves open, wide
behind. Antenna (fig. 112) five-segmented; scape (2:1) slightly more than one
and a half times as long as the pedicel (2:1), the first flagellar segment short,
about one fifth the length of the second (10:7), which is three quarters the
length of the apical segment (8 : 3). Mouthparts, figs. 111, 116.
Thorax, fig. 113. Pronotum more than two and a half times as long as wide
anteriorly, and twice as long as its maximum width. Mesonotum wider than long
{5:4}. Metanotum not quite as wide as the mesonotum, and incompletely
separated from the propodeum, which is nearly as long as wide. Fore leg (fig.
109): femur more than twice as long as the tibia. Tibial armature consisting of
four dorso-apical teeth, one of which is very small, and two ventral teeth. Tarsal
segments in ratio 5 : 6. Mid leg slender, the coxa nearly as long as trochanter and
femur combined, and as long as the clavate tibia. The tibia apically armed with a
series of sharp teeth (fig. 107). Tarsus pentamerous, the first segment slightly
longer than the fifth [14 : 13}, the intermediates subequal, not quite half as long
as the fifth {5 :12]. Hind leg (fig. 106) rather long, the femur as long as coxa
and trochanter combined; all these segments slender. Tibia nearly two-thirds the
length of the femur, with a dorsal hyaline ridge. Ventro-apical armature and
pubescence as in the figure. Tarsus dilated, the first segment as long as the tibia,
the second half as long; third to fifth segments subequal, two-thirds the length
of the second. Pubescence half as long as the width of the segments.
Gaster. Cerci of the tenth urite with two claws, aedeagus slightly dilated sub-
apically and apically (fig. 114).
Length, 1.1 mm. Colour uniformly yellow-brown.
Female. Antenna consisting of eleven segments, of which the apical two form
a loose club. Scape approximately twice as long as the pedicel, appendage of the
third segment rather short and robust. Fifth segment three quarters the length of
the seventh, the eighth segment slightly longer than the seventh, ninth to eleventh
segments gradually diminishing in size. The funicular segments bear two irregular
rows of long sensilla. Maxilla without a bacilliform process, but with a long,
subapical hair. Mandible (fig. 110) with four ventral ridges, its appendage with
five ridges.
Thorax. A few hairs occur above and beneath the propodeal spiracle. Fore
wing (2:1), 1.4 mm long. Submarginal, marginal, stigmal, and postmarginal
veins in ratio 20 : 7 : 7 : 11, the submarginal vein with three pustules, the stigmal
vein with four pustules. Hind wing (4:1), 0.8 mm long. Coxa of the fore leg
(fig. 108) two-thirds the length of the femur, the tibia not quite half as long as
the femur. Tibial armature consisting of four dorsal teeth. Tarsal segments ap-
proximately in ratio 11:7:5:4:5. Mid leg slender, the subclavate tibia not
quite as long as femur and trochanter combined. First tarsal segment twice as long
as the second, second to fifth segments subequal. Hind leg (fig. 115): length of
the coxa two-thirds of that of the femur, which is not nearly twice as long as the
J. T. WieBEs: Indo-Australian Ceratosolen 37
Figs. 130—143, Ceratosolen vechti, Java (139, specimen from Sumatra), 130, male labium
and maxillae, 131-133, female, 131, antenna, 132, labium and maxillae, 133, hind leg,
134, male, 135, female mandible, 136—142, male, 136, fifth tarsal segment of hind leg,
137, fore leg, 138, apex of abdomen, 139, cercus of tenth urite and paramere, specimen from
Sumatra, 140, hind leg, 141, antenna, 142, mandible, 143, female fore leg. Figs. 134, 140,
DESSEN 31437505 ZIA X 155: 138 139 14210, 2155 130; 136
DCP)
38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
tibia [12 : 7}. The antaxial tibial tooth is bidentate; the axial tooth, usually found
in the species of Ceratosolen, is lacking in all specimens studied. Tarsus pubescent,
especially along the plantar edge, the segments with heavy ventro-apical spines.
Tarsal segments approximately in ratio 25:5:5:4:5.
Gaster. The ovipositor distinctly projecting beyond the apex of the gaster.
Length, 1.3—1.4 mm.
Remark.
Due to the fact that the samples have been desiccated, this species could not be
described and figured in the usual detail. Notably the chaetotaxy must for the
greater part remain undescribed. Yet, C. gracilis is easily recognizable by its size,
by the relatively long hind legs of the male, and by the absence of a bacilliform
process from the female maxilla.
Ceratosolen hooglandi sp. n. (figs. 117—129)
Material.
Series 9, 4, ex Ficus bernaysii King (det. E. J. H. CORNER), Mawan Village
(Gogol Valley, Madang distr., Terr. New Guinea), leg. R. D. HOOGLAND,
16.VI.1955, no. 4890; coll. no. 362; & (holotype), slide 362a, :? (allotype),
362c, 4, 9 (paratypes), 362b, d.
Series ©, 4, ex Ficus bernaysi King (det. E. J. H. CORNER), Lae (Terr. New
Guinea), Busu River, leg. E. J. H. CORNER, 8.IX.1960, NGF 12471; coll. no. 517.
Description.
Male. Head (fig. 127) more than twice as long as wide anteriorly, and one
and a half times as long as its maximum width. Pubescence short and sparse.
Epistomal margin with four short hairs. Eyes absent. Antennal grooves open, not
very wide behind. Antenna (fig. 121) consisting of five segments. Pedicel (5 : 3)
more than half as long as the scape (8 : 5); the first funicular segment (5 : 8)
half as long as the pedicel, the second flagellar segment (5 : 4) half as long as
the first; the apical segment (3 :1) more than twice as long as the second. Man-
dible, fig. 122. Maxilla and labium, fig. 119; maxilla with one lateral hair.
Thorax, fig. 127. Pronotum nearly twice as long as wide anteriorly; the maxi-
mum width four-fifths of the length. Length of the mesonotum three quarters of
the width. Metanotum incompletely separated from the propodeum, slightly nar-
rower than the mesonotum. Tibia of the fore leg (fig. 120) not nearly half as
long as the femur, with three large dorsal and three ventro-apical teeth. Second
tarsal segment slightly longer than the first. Coxa of the mid leg as long as the
femur, tibia nearly as long as femur and trochanter combined. Tibia with only
one inconspicuous apical tooth. Tarsal segments in ratio 3:1:1:1:3. Hind leg,
fig. 118. Coxa and tibia shorter than the femur [7 : 8}. Tibia one third longer
than the first tarsal segment, which is as long as third and fourth segments
combined. Second, third, and fifth segments subequal, fourth smaller. Tarsal
pubescence distinctly shorter than half the width of the segments; dilatation not
very wide.
J. T. WieBESs: Indo-Australian Ceratosolen 39
Gaster. Cerci of the tenth urite with three short claws, aedeagus not dilated at
apex (fig. 126).
Length, 1.4 mm. Colour uniformly yellowish brown.
Female. Head longer than wide between the external margins of the eyes
{10 : 9}. Longitudinal diameter of the eye little shorter than the cheek. Pubescence
rather long. Antenna (fig. 124): the scape relatively small, about as long as third
to fifth segments combined. Pedicel slender, and little more than one third the
length of the scape. Fourth segment very small, the fifth more than twice as
long. Sixth segment more than one and a half times as long as the fifth, seventh
to tenth segments gradually diminishing in size; the eleventh as long as the fifth.
Funicular segments with two rows of long narrow sensilla. Mandible (fig. 125)
with four ventral ridges; the appendage shorter than the mandible, with five ridges.
Maxilla (fig. 117) with a long bacilliform process, which is nearly half as long as
the maxilla. Maxilla as well as its process with a long hair; labium without hairs.
Thorax slender, its parts with many long hairs. Scutum as long as wide anteriorly,
with approximately ten hairs on each side. Scutellum little longer than wide
anteriorly, with lateral rows of fifteen to twenty hairs. Metanotum with five hairs
on each side of the mid-line. Propodeum wide; little groups of hairs are situated
above and beneath the spiracle. Forewing (7:3), 1.4 mm long. Submarginal,
marginal, stigmal, and postmarginal veins approximately in ratio 30:9 : 11 : 17.
Submarginal vein with three pustules, stigmal vein (fig. 123) with four. Hind
wing (5:1), 0.8 mm long. Fore leg, fig. 128. Coxa nearly as long as the femur,
tibia half as long. Tibia with four teeth along the dorsal edge, ventral edge with
one tooth. Tarsal segments in ratio 3:2:2:2:3. Mid leg slender, tibia shorter
than trochanter and femur combined [7 : 8}; tarsal segments in ratio 12:5 : 4:
4:7. Coxa and femur of the hind leg (fig. 129) subequal in length, the tibia
shorter {8 : 11}. Tibia with the usual two apical teeth, the one bidentate, and the
other unidentate. First tarsal segment as long as the tibia, and nearly thrice as
long as the fifth tarsal segment. Intermediate segments shorter than the fifth
{2:3}. Ventral edge of the tarsus fimbriated.
Gaster. Ovipositor scarcely projecting beyond the apex of the gaster.
Length, 1.4 mm. Colour dark brown, with lighter ventral surface and legs.
Nervures of wings yellowish brown.
Remark.
C. hooglandi is recognizable by the long and slender tibia and metatarsus of
the male hind leg, and by the relatively long bacilliform process of the female
maxilla.
It is a pleasure to name the species after its collector, Dr. R. D. HOOGLAND.
Ceratosolen vechti sp. n. (figs. 130—143; Pl. 1)
Material.
Series 9, 4, ex Ficus lepicarpa Bl.1) (det. E. J. H. CORNER), Tjibodas (Java),
1) Discussion of host record, see p. 98.
_40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
AN
u
Al N
n |
Figs. 144— 151, Ceratosolen nugatorius, Kelantan, male, 144, hind leg, 145, fore leg, 146,
maxilla, 147, antenna, 148, mandible, 149, mid leg (coxa omitted), 150, apex of abdomen,
151, male, 152—153, C. bisulcatus, Java, 152, male labium and maxillae, 153, female labium
and maxillae. Figs. 151, X 60; 144, 145, 149, X 95; 147, X 185; 148, 150, 152, 153,
NE 2558 146, II
J. T. Wirprs: Indo-Australian Ceratosolen 41
along stream, 1500 m alt, leg. J. H. DE GUNST, 16.X1.1954; coll. no. 98; 4
(holotype), slide 98a, ® (allotype), 98d, 4, 9 (paratypes), 98b, c, e, f.
Six 9,5 &, ex Ficus lepicarpa Bl, Tjibodas (Java), Mt. Gedeh, 1400 m alt,
leg. J. H. DE GUNST, 16.X1.1954; coll. no. 6.
Fragments ©, ex Ficus lepicarpa Bl, Tjibodas (Java), leg. J. H. DE Gunsr,
5.XI.1954; coll. no. 70.
Eleven 9, ex Ficus lepicarpa Bl, Tjibodas (Java), leg. J. H. DE Gunsr,
11.11.1955; coll. no. 300.
Four 9, series 4, ex Ficus lepicarpa BI. (det. E. J. H. CORNER), Pajakumbuh
(Sumatra), Mt. Sago, leg. W. MEIJER, 14.V.1955, no. 3334; coll. no. 371; &,
slide 371a.
? Six @, ex Ficus lepicarpa BI. (det. E. J. H. CORNER), Kundasan (N. Bor-
neo), 4000 ft. alt, leg. E. J. H. CORNER, 6.IX.1961; coll. no. 624.
DESC PD EO
Male. Head (fig. 134) more than twice as long as wide anteriorly, its maximum
width three quarters of the length. Epistomal margin with four short hairs.
Pubescence short. Eyes small. Antennal grooves open, not very wide behind.
Antenna (fig. 141) five-segmented; pedicel (5:3) half as long as the scape
(2:1). First flagellar segment (2 : 3) small, the apical (11 : 3) somewhat longer
than the scape, and more than one and a half times as long as the subapical seg-
ment (7 : 4). Mandible, fig. 142. Labium and maxillae, fig. 130; maxilla with
two hairs at mid length.
Thorax, fig. 134. Pronotum twice as long as wide anteriorly, the maximum
width nearly three quarters of the length. Mesonotum with rounded edges, one
and a half times as wide as long. Metanotum incompletely separated from the
propodeum, approximately as wide as the mesonotum; propodeum narrower, and
more than twice as long as the metanotum. Femur of the fore leg (fig. 137) more
than twice as long as the tibia, which bears four dorsal and three ventro-apical
teeth. Tarsus bimerous, the second segment is the longer. Coxa of the mid leg
nearly as long as the femur [5 : 6}, the trochanter small. Tibia longer than the fe-
mur [7 : 6}, apically with three teeth, the dorsal one of which is particularly long.
Tarsal segments in ratio 3:2:2:2:5. Hind leg, fig. 140; coxa, femur, and
tibia about equal in length {10 : 11 :9}, the tibia as long as the first and second
tarsal segments combined. First tarsal segment one and a half times as long as
the fifth (fig. 136), the fourth as long as the fifth, second and third segments
smaller. Tarsus dilated, with pubescence approximately as long as one third the
width of the segments.
Gaster. Cerci of tenth urite with two claws, aedeagus not dilated at apex
(fig. 138). Some specimens from Sumatra differ in having the cerci with three
or four claws, and the parameres with a subapical hair (fig. 139).
Length, 1.7 mm. Colour yellowish, with darker hind tarsi in some specimens, or
only the distal segments brown.
Female. Head as long as wide across the compound eyes. Longitudinal diameter
of the eye about as long as the cheek. Pubescence short. Antenna (fig. 131) with
42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
the scape as long as third to sixth segments together; the pedicel half as long as
the scape, the fourth segment small, the fifth approximately twice as long. Seg-
ments six to ten subequal, the eleventh smaller. Two apical segments shaped so as to
form together a club. Sensilla of the funicle numerous, rather wide, mainly situated
in the distal parts of the segments. Mandible (fig. 135) with six ventral ridges,
the appendage but little shorter than the mandible, with six ridges. Maxilla (fig.
132) with a bacilliform process, which is one third the length of the maxilla, and
a subapical hair; labium and the process with an apical hair.
Thorax. Pronotum wide and bristly. Scutum shorter than wide anteriorly [2 : 3},
scutellum nearly one and a half times as long as wide anteriorly. Scutum with
three hairs, scutellum with seven to ten hairs along each lateral margin. Metanotum
with eight hairs on each side of the mid line. Propodeum with little groups of
hairs above and beneath the spiracle. Fore wing (2:1), 2.0 mm long. Sub-
marginal, marginal, stigmal, and postmarginal veins in ratio 42 : 15 : 13 :23. Sub-
marginal vein with two or three pustules, stigmal vein with four (or sometimes
five). Hind wing (5 : 1), 1.2 mm long. Coxa of the fore leg (fig. 143) two-thirds
the length of the femur; the tibia not half as long as the femur, with four dorsal
and one ventro-apical teeth. First tarsal segment a little longer than the fifth
[5 : 4}, intermediates subequal, half as long as the fifth. Coxa and trochanter of the
mid leg short, the tibia longer than the femur {4 : 3}, tarsal segments in ratio 7:3:
3 :2:4. Tibia and tarsus rather bristly. Coxa of the hind leg (fig. 133) nearly as
long as the femur, trochanter small. Tibia with the usual two apical teeth, more
than half as long as the femur. Tarsal segments in ratio 11:4:3:3:6, with
fimbriated ventral edges.
Gaster. Ovipositor barely projecting beyond the apex of the gaster.
Length, 2.2 mm. Colour dark brown, legs and ventral surface lighter. Nervures.
of the wings light brown.
Remark.
This species is remarkable by the robust hind legs of the males, and by the
wide sensilla of the female antenna.
It is a pleasure to name this species after Dr. J. VAN DER VECHT, Professor of
Systematic Zoology at Groningen University, Curator of Hymenoptera, Rijks-
museum van Natuurlijke Historie, Leiden, who recognized it as new when studying
the samples from Java.
Ceratosolen nugatorius Grandi (figs. 144—151)
Ceratosolen nugatorius Grandi, 1952b, pp. 55—57, figs. V—VI [descr. 9, ex Ficus spec.
Kuala Lumpur (Malaya), IX.1948].
Material.
Three 9, Kuala Lumpur (Malaya), ex coll. GRANDI; coll. ML, no. 510.
Series ©, 4, ex Ficus obpyramidata King (det. E. J. H. CORNER), Batu Papan
(Kelantan, Malaya), leg. E. J. H. Corner, Sing. F. no. 29526; coll. no. 368;
3 (allotype), slide 368a, 9, 4, 368b-e.
J. T. WIEBES : Indo-Australian Ceratosolen 43
Figs. 154—166, Ceratosolen dentifer, New Guinea, 154—156, female, 154, antenna, 155,
mandible, 156, detail of hind leg, 157—159, male, 157, hind leg, 158, detail of hind leg,
159, apex of mid tibia, 160, detail of female fore leg, 161—166, male, 161, antenna, 162,
maxilla, 163, detail of fore leg, 164, cercus of tenth urite and paramere, 165, mandible,
166Mmale Fes MMS TIGENO: IS AMIS GMS AGS: 155, 159—161, 165, X 155;
162, 164, X 365
44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Two 2,7 4, ex Ficus obpyramidata King (det. E. J. H. CORNER), Kemanan
(Trengganu, Malaya), leg. E. J. H. CORNER, Sing. F. no. 25843; coll. no. 365;
&, slide 365a.
Description — Additional note.
I consider the specimens from Ficus obpyramidata conspecific with C. nugatorius
Grandi, and describe the hitherto unknown male.
Male. Head (fig. 151) twice as long as wide anteriorly, its maximum width
more than two-thirds the length. Pubescence short. Epistomal margin with four
short hairs. Eyes very small. Antennal grooves open, narrow behind. Antenna
(fig. 147) five-segmented; the pedicel (2:1) two-thirds the length of the scape
(9 : 5), the first flagellar segment (7 : 6) three quarters the length of the second
(2:1), the apical segment (7:2) nearly twice as long as the subapical one.
Mandible, fig. 148. Maxilla (fig. 146) with two lateral hairs.
Thorax, fig. 151. Pronotum one and a half times as long as wide anteriorly, and
but little longer than its maximum width. Maximum width of the mesonotum
approximately five-thirds of the length. Metanotum almost completely separated
from the propodeum, propodeum subquadrate. Fore leg (fig. 145): the tibia ap-
proximately half as long as the femur, with four dorsal and two ventro-apical
teeth. Tarsal segments in ratio 4 : 3. Coxa and femur of the mid leg subequal in
length, the tibia much longer {4 : 3}, with four sharp, apical teeth (fig. 149).
Tarsal segments in ratio 11:7:7:8:17. Hind leg, fig. 144. Coxa little shorter
than the femur, and nearly one and a half times as long as the tibia. Tibia with
a large motile spur on the disk. First tarsal segment little shorter than the tibia,
but longer than the fifth, intermediate segments shorter, subequal. Tarsus dilated,
the pubescence nearly as long as half the width of the segments.
Gaster. Cerci of the tenth urite with four short claws (two of which are small
and sometimes difficult to be observed), the aedeagus dilated (fig. 150).
Length, 1.3 mm. Colour reddish brown. Head darker than the other parts of
the body, hind tarsi light brown.
Remark.
The peculiar spur of the male hind leg, the short and robust male thorax, and
the very few sensilla of the female antenna, distinguish this species from its
congeners.
Ceratosolen dentifer sp. n. (figs. 154—166)
Material.
Series 9, 4, ex Ficus hispidioides S. Moore var. succosa Corner (det. E. J. H.
CORNER), Lae (Terr. New Guinea), leg. E. J. H. CORNER, X.1960; coll. no. 524;
& (holotype), slide 524a, 9 (allotype), 524c, 9, & (paratypes), 524b, d.
Series 9, 4, ex Ficus hispidioides S. Moore var. succosa Corner (det. E. J. H.
CORNER), Lae (Terr. New Guinea), leg. E. il H. CORNER, 5.IX.1960; coll. no.
S22 sli dei522a 20255225!
J. T. WieBEs: Indo-Australian Ceratosolen 45
Description.
Male. Head (fig. 166) twice as long as wide anteriorly, the maximum width
two-thirds of the length. Pubescence short. Eyes small. Antennal grooves open, not
very wide behind. Antenna (fig. 161) five-segmented; the scape (2:1) more
than twice as long as the pedicel (5:4), the first flagellar segment short, the
second approximately twice as long as the first, and half as long as the pedicel;
the apical segment (5:1) thrice as long as the pedicel. Mandible, fig. 165.
Maxilla (fig. 162) with three lateral hairs, labium without long hairs.
Thorax, fig. 166. Pronotum not quite twice as long as wide anteriorly, the
posterior width slightly smaller than the length. Mesonotum nearly twice as
wide as long. Metanotum short, incompletely separated from the large propodeum,
which is nearly as long as wide [8 :9}. Fore leg, fig. 163. Femur twice as long
as the tibia, which bears four dorso-apical teeth and three ventral teeth. Tarsal
segments in ratio 4 : 5. Coxa, femur, and tibia of the mid leg subequal in length,
the tibia (fig. 159) with seven apical teeth. Tarsal segments in ratio 5: 2:3 :3 :8.
Hind leg (figs. 157—158): coxa smaller than the femur, the tibia about half as
long. Apex of the tibia with a motile spur (fig. 158). Tarsus dilated, the first seg-
ment as long as the tibia, the second segment half as long, third and fourth seg-
ments slightly smaller than the second. Fifth segment two-thirds the length of the
first. The segments gradually widening distally; pubescence approximately half as
long as the width of the segments.
Gaster. Cercus of the tenth urite (fig. 164) with three or four claws.
Length, 1.6 mm; very small specimens measure 1.2—1.3 mm. Colour uniformly
yellow-brown, head a trifle darker.
Female. Head as long as wide across the compound eyes, the cheeks as long as
the eyes. Pubescence sparce, not very long. Antenna, fig. 154. Scape two and a
half times as long as the pedicel; the pedicel with approximately fifty-five small
spines on the axial surface. Fifth segment, with six long sensilla, shorter than the
sixth, and about as long as the tenth. Seventh to tenth segments gradually
diminishing in size, the eleventh, which forms a loose club with the tenth, one
and a half times as long as the tenth. Funicular segments with regular rows —
one per segment — of seven to nine long sensilla, the apical segment moreover
with two circular pits. Mandible (fig. 155) with six ventral ridges, its appendage
with five ridges. Maxilla with a subapical hair, and a bacilliform process, which
is less than half as long as the maxilla [9 : 22}, and bears a long apical hair.
Labium without long hairs.
Thorax. Pronotum with sparce, long pubescence. Scutum one and a half times
as long as wide maximally, and thrice as long as wide posteriorly, with seven to
nine hairs along the lateral margins. Scutellum as long as wide posteriorly, nar-
rower anteriorly, with eight to nine hairs along the lateral edges. Metanotum with
ten hairs anteriorly, and two posteriorly, on each side of the mid line. Propodeum
wide, with two hairs above, and a group of approximately ten hairs below the
spiracle. Fore wing (2:1), 1.4 mm long. Submarginal, marginal, stigmal, and
postmarginal veins approximately in ratio 14:5 :5 : 12. Submarginal vein with
three pustules, stigmal vein with four. Hind wing (9:2), 1.1 mm long. Femur
46 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Figs. 167—178, Ceratosolen moderatus, Borneo, 167, female antenna, 168, male antenna,
169, detail of female hind leg, 170, detail of male fore leg, 171, detail of female fore leg,
172—176, male, 172, labium and maxilla, 173, male, 174, hind leg, 175, detail of hind
leg, 176, detail of hind leg, same specimen as 175, axial aspect of other leg, 177, female
mandible, 178, male cercus of tenth urite and paramere. Figs. 173—176, X 60; 169—171,
X 140; 167, 168, 177, X 185; 172, 178, X 440
J. T. Wieses: Indo-Australian Ceratosolen 47
of the fore leg (fig. 160), with long pubescence along the dorsal and ventral
margins, nearly twice as long as the coxa, and more than thrice as long as the
tibia. Tibia with four dorsal and one ventro-apical teeth. Tarsus pentamerous,
with rows of spines; segments in ratio 19:6:6:4:9. Mid leg slender; the tibia
nearly as long as trochanter and femur combined, tarsal segments in ratio 16 :9 :
8 : 6:9. Tibia and tarsus with long pubescence, tarsal segments with apical spines.
Hind leg, fig. 156. Coxa smaller than the femur, the tibia more than half as long
[13 : 25}. Femur heavily pubescent. Antaxial tibial spur bidentate, the axial one
curved. Tarsus with long pubescence and apical spines; segments in ratio 16:8:
GANT:
Gaster. Ovipositor slightly protruding behind the gaster.
Length, 2.2—2.4 mm (very small specimens measure 1.8 mm). Colour: head
and dorsal surface of thorax dark brown, gaster lighter.
Extremities yellow-Erown. Nervures of the wings brown.
Remark.
The tibial spur of the male hind leg is a common character of C. dentifer and
C. nugatorius; they are, however, distinguishable by the male and female mouth-
parts, the relative proportions of the male thorax, etc.
Ceratosolen moderatus sp. n. (figs. 167—178)
Material.
Series 9, 4, ex Ficus moderata Corner (det. E. J. H. CORNER), Kundasan
(N. Borneo), 3500 ft. alt., leg. E. J. H. CORNER, 31.VIII.1961, RSNB no. 2594;
coll. nos. 628, 647, 649, 651; 4 (holotype), slide 628a, 9 (allotype), 628d,
3, 9 (paratypes), 628b, c, e.
Description:
Male. Length of the head (fig. 173) one and a half times the maximum
width, and more than twice the anterior width. Eyes small. Antennal grooves
open, not very wide behind. Antenna (fig. 168) five-segmented, the scape (3 : 2)
one and a half times as long as the pedicel (2:1); the first flagellar
segment annuliform, the second nearly as long as the pedicel, and more than half
as long as the apical segment (5 : 2). Maxilla (fig. 172) with two hairs, the sub-
apical one of which is absent in some specimens; labium without long hairs.
Mandible as in the preceding species.
Thorax, fig. 173. Pronotum slightly shorter than wide posteriorly, and twice as
long as wide anteriorly. Mesonotum approximately twice as wide as long. Meta-
notum short, incompletely separated from the propodeum, which is distinctly
wider than long (3 : 2). Femur of the fore leg (fig. 170) twice as long as the
tibia, with few hairs. Tibia with three dorsal and three ventro-apical teeth. Tarsus
bimerous, the segments approximately equal in length; axially with a few stout
hairs. Mid leg slender, the femur shorter than the tibia [6 : 7}, the coxa shorter
than the femur {5 : 6}, but wider. Tibia with three apical teeth. Tarsal segments
approximately in ratio 10 : 5 :4:5 :10. Oligomery, in casu, incomplete separation
48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
of second, third, and fourth segments, occurs sporadically. Hind leg, figs. 174—
176. Coxa and femur subequal in length, the tibia shorter. Tibia with a
bidentate apical tooth at the antaxial edge and a ventral spur. Tarsus dilated,
segments (in the holotype) in ratio 25 : 16 : 11 : 15 :16; the pubescence as long
as one third the width of the segments. There is a considerable variation in the
relative proportions of the tarsal segments, which may be wider than shown in fig.
174, but nearly always the second and fifth segments are subequal, and the first
distinctly larger. The third and fourth segments are nearly as long as the second,
or longer, or as drawn in fig. 174. Moreover, some of the specimens with large
hind feet show oligomery: the first and second, second and third, or fourth and
fifth tarsal segments being incompletely separated (figs. 175, 176).
Gaster. Cerci of the tenth urite large, with five small claws (fig. 178).
Length, 1.2—1.3 mm. Colour uniformly light yellow-brown.
Female. Head as long as wide across the compound eyes. Longitudinal diameter
of the eye as long as the cheek. Pubescence not very long. Antenna (fig. 167):
scape large, four times as long as the pedicel, which bears approximately forty
axial spines. Third segment not very long, the fourth small. Fifth segment with
ten long sensilla. Sixth to tenth segments subequal, distinctly longer than the
fifth, the eleventh shorter, about as long as the fifth. Eleventh segment with six
oblong and two circular sensilla, other segments with two rows of six to nine
sensilla. Maxilla with one subapical hair, and with a bacilliform process, which
is half as long as the maxilla, and bears one apical hair. Mandible (fig. 177)
with six large ventral ridges, and five ridges on the appendage.
Thorax. Pronotum with sparse, but long pubescence. Scutum as long as wide
maximally, the posterior width slightly more than two-thirds the length. Seven
to eight hairs occur on each lateral side. Anterior width of the scutellum four-
fifths, posterior width six-sevenths of the length, lateral margins with about ten
hairs. Metanotum with eight hairs on each side of the mid line; propodeum with
three hairs above, and six hairs below the spiracles. Fore wing (7 :3), 1.65 mm
long. Submarginal, marginal, stigmal, and postmarginal veins in ratio 14:4:5:
11; submarginal vein with two pustules, stigmal vein with four. Hind wing
(5:1), 0.95 mm long. Fore leg (fig. 171): length of the coxa three-fifths, length
of the tibia two-fifths of that of femur and trochanter combined. Tibia with four
dorsal and one ventral teeth. Tarsal segments in ratio 40 : 13 :13 :14 : 20. Mid
leg slender, the tibia nearly as long as femur and trochanter combined, with long
hairs. Tarsus with long pubescence, segments in ratio 15 :5:5:5:6. Coxa of
the hind leg (fig. 169) slightly shorter than the femur; the tibia two-thirds of
the femur, with an antaxial bidentate, and an axial unidentate tooth at the ventro-
apical edge. Tarsal segments wide, ratio: 30:9:6:5:9.
Gaster. Ovipositor slightly projecting beyond the apex of the gaster.
Length, 1.7—1.8 mm. Colour: head and dorsal surface of body grey-brown,
ventral surface and extremities lighter. Nervures of the wings brown.
Remark.
C. moderatus is not much larger than C. crassitarsus, but it is easily recognized
by the relative proportions of the male antenna and hind leg, the heteromery of
J. T. Wreses: Indo-Australian Ceratosolen 49
Figs. 179—191, Ceratosolen praestans, New Britain, 179, female antenna, 180, male, 181,
female fore leg, 182, male labium and maxillae, 183, female mandible, 184—186, male,
184, cercus of tenth urite, 185, mid leg, 186, mandible, 187, detail of female hind leg, 188,
detail of male fore leg, 189, male hind leg, 190, female maxilla, 191, male antenna. Figs.
180, 189, X 40; 179, 181, 185, 187, 188, X 90; 183, 186, 190, 191, X 115; 182, X 160;
184, X 200
the male hind tarsus, and the presence of five claws on the cerci of the tenth urite.
The females have hyaline wings without striae.
Ceratosolen praestans sp. n. (figs. 179—191)
Material.
Series 9, &, ex Ficus praestans Corner (det. E. J. H. CORNER), Keravat (New
Britain), leg. E. J. H. CORNER, X.1960; coll. no. 533; & (holotype), slide 533a,
Q (allotype), 533c, &, 9 (paratypes), 533b, d.
50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
DES EFD Ll OE
Male. Head (fig. 180) nearly twice as long as wide anteriorly, and one and a
half times as long as its maximum width. Epistomal margin prominent, with six
hairs. Dorsal and ventral surfaces with sparse, thin hairs. Eyes wanting. Antennal
grooves open, not very wide behind. Antenna (fig. 191) five-segmented. Scape
(5 : 3) nearly twice as long as the pedicel (3 : 2); the first flagellar segment short,
the second (5 : 4) slightly shorter than the pedicel, the apical segment (3 : 1)
more than twice as long. Mandible, fig. 186. Labium and maxillae, fig. 182;
labium small, maxilla large, with one apical hair.
Thorax, fig. 180. Pronotum not quite twice as long as wide anteriorly, and
distinctly longer than wide posteriorly [7 : 6}. Mesonotum with rounded edges,
its length two-thirds of the maximum width. Metanotum short, incompletely
separated from the propodeum. Propodeum slightly tapering behind, its length
three quarters of the maximum width. Femur of the fore leg (fig. 188) twice as
long as the tibia. Tibia with four dorsal and three ventro-apical teeth. Tarsus con-
sisting of two segments, the first of which is the longer [6:5]. Mid leg (fig.
185): the coxa a little shorter than the femur; the tibia as long as coxa and
trochanter combined. Tibia with five apical teeth: three ventrals and two dorsals.
Tarsus pentamerous, the segments in ratio 11 : 5 : 5 : 4:15. Hind leg (fig. 189):
the coxa slightly shorter than the femur, but longer than the tibia. Dorsal edge of
the coxa, axial surfaces of femur and tibia, and dorsal edge of the tibia pubescent.
Tibia with three ventral apical teeth. Tarsus dilated, with pubescence approximate-
ly as long as half the width of the segments. Segments in ratio 29:12:12 :13:
16.
Gaster. Aedeagus dilated apically. Cerci of the tenth urite (fig. 184) with seven
claws, parameres tapering distally.
Length, 1.6 mm. Colour uniformly yellow-brown.
Female. Head slightly longer than wide between the external margins of the
eyes. Longitudinal diameter of the eye a little longer than the cheek {6 : 5]. An-
tenna, fig. 179. Scape as long as second to fifth segments combined. Pedicel not
quite half as long as the scape, with approximately forty-five slender spines. Fifth
segment with two rows of four sensilla, and two apical sensilla which are oriented
perpendicularly to the others; the sixth segment with four sensilla in the basal
row, and seven apical sensilla, seventh to tenth segments with two rows of six
to eight sensilla; the apical segment with five sensilla. Mandible (fig. 183), and
its appendage, with five ventral ridges. Labium without hairs. Maxilla (fig. 190)
with one subapical hair, and a bacilliform process. The bacilliform process, which
is about one fifth the length of the maxilla, seems to be very delicate, and it is,
more often than not, broken off close to its base, or even entirely absent.
Thorax. Pronotum wide, with many hairs. Scutum nearly as long as wide an-
teriorly, scutellum one and a half times as long as wide anteriorly. Metanotum
nearly as wide as scutum and scapulae together, with ten hairs on each side of
the mid line. Propodeum wider, with six hairs above, and twelve hairs beneath
the spiracular peritremata. Fore wing (2 : 1), 2.2 mm long. Submarginal, marginal
stigmal, and postmarginal veins in ratio 17 :7:5:13. Submarginal vein with
J. T. WieBEs: Indo-Australian Ceratosolen 51
three pustules, stigmal vein with four. Hind wing (4: 1), 1.3 mm long. Coxa of
fore leg (fig. 181) not quite two-thirds the length of the femur, tibia much
shorter than the femur [2 : 5}, with four teeth at the antaxial, dorso-apical edge,
and one ventral tooth. Tarsal segments in ratio 16:4:5:6:9; first segment
with ventral spines, other segments with hairs and apical spines. Coxa and tro-
chanter of the mid leg short; the femur four times as long as the trochanter, the
tibia nearly as long as femur and trochanter combined. Tarsal segments in ratio
10:4:4:4:5. Hind leg, fig. 187. Coxa slightly smaller than the femur, the
tibia half as long as the femur. Tibia with one ventral axial tooth, and a bidentate
antaxial tooth. Tarsus pubescent, especially along the plantar edge; segments in
Mio 2 slee BOO) Bil
Gaster. Ovipositor scarcely projecting beyond the apex of the gaster.
Length, 2.4—2.6 mm. Colour dark brown, legs and antennae lighter. Wings
with dark striae radiating from the stigma.
Remark.
This species is similar to some other species in which the hind feet of the males
are dilated. It is distinguished by the male genitalia (cerci with seven claws) and
by the short bacilliform process of the female maxilla.
Ceratosolen iodotrichae sp. n. (figs. 192—203)
Material.
Series 9, &, ex Ficus iodotricha Diels (det. E. J. H. CORNER), Wau (Terr.
New Guinea), Edie Creek, leg. E. J. H. CORNER, 19.IX.1960, NGF no. 12498;
coll. no. 535; 4 (holotype), slide 535a, 9 (allotype), 535c, 9, 4 (paratypes),
55 Dd mer
Descriptions
Male. Head (fig. 198) twice as long as wide anteriorly, with almost parallel
sides. Pubescence short and sparse. Eyes wanting. Antennal grooves open, not
very wide behind. Antenna (fig. 196) five-segmented; the scape (2:1) not
quite twice as long as the pedicel (3 :2); the first flagellar segment (1 :2)
slightly shorter than the second (2:3), the apical segment (4:1) six times as
long as the second. Mandible, fig. 200. Maxilla (fig. 199) with two lateral hairs;
labium without long hairs.
Thorax, fig. 198. Pronotum nearly one and a half times as long as wide
posteriorly, much narrower anteriorly. Maximum width of the mesonotum
distinctly longer than the length {7 : 5}. Metanotum incompletely separated from
the propodeum, as wide as the mesonotum. Propodeum approximately as long as
wide, its posterior margin nearly straight. Fore leg (fig. 203): the tibia, without
the apical teeth, half as long as the femur. Tibia with four dorsal and four ventro-
apical teeth. Tarsus bimerous, distal segment slightly longer than the proximal one.
Femur and tibia of the mid leg subequal in length, slightly shorter than coxa and
trochanter combined. Tibia with seven apical teeth: six at the ventral margin, and
one bidentate, dorsal tooth. Tarsus pentamerous, segments in ratio 4:2:2:2:5.
52
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Figs. 192—203, Ceratosolen iodotrichae, New Guinea, 192, male hind leg, 193—195, female,
193, mandible, 194, maxilla, 195, antenna, 196—200, male, 196, antenna, 197, cercus of
tenth urite, 198, male, 199, maxilla, 200, mandible, 201, detail of female fore leg, 202,
detail of female hind leg, 203, detail of male fore leg. Figs. 192, 198, X 50; 195, 202, 203,
NOELS wie, SHY, IGG, 0020 MS IMO Oe) S< BED)
J. T. WIEBES : Indo-Australian Ceratosolen 53
Hind leg (fig. 192) much like that of C. hoogland, but more robust; the tibia
with one small apical tooth. Tarsal segments in ratio 14:10 :9 :9 : 11; dilated,
pubescence approximately as long as one third the width of the segments.
Gaster. Cerci of the tenth urite (fig. 197) with four or five claws.
Length, 1.7—1.8 mm. Colour uniformly yellowish brown.
Female. Head as long as wide across the compound eyes. Eyes protruding, about
as long as the cheek. Pubescence sparse, but long. Antenna (fig. 195); pedicel
not half as long as the scape, with approximately forty short spines axially. Fifth
segment twice as long as the short fourth, sixth to ninth subequal, longer than
the fifth, tenth and eleventh smaller, together forming a loose club. Funicular
segments with rather long sensilla: the fifth with twelve, sixth to tenth with two
irregular rows of six to eight sensilla. Apical segment with six oblong and two
circular sensilla. Mandible (fig. 193) with six ventral ridges, its appendage with
four. Maxilla (fig. 194) with one, or sometimes two, subapical hairs, and a
bacilliform process, which is, without the apical hair, between one third and one
half the length of the maxilla. Labium bare.
Thorax. Scutum approximately one and a half times as long as wide posteriorly,
much wider anteriorly; seven hairs occur along each lateral margin. Scutellum little
wider behind than in front, its length four-thirds of the posterior width. Metano-
tum short, with eight hairs on each side of the mid line. Propodeum with six
hairs above, and approximately twice as many behind the spiracle. Fore wing
(8:3), 2.6 mm long. Submarginal, marginal, stigmal, and postmarginal veins in
ratio 24:5: 7:13; submarginal vein with three pustules, stigmal vein with four.
Hind wing (4:1), 1.4 mm long. Femur of the fore leg (fig. 201) nearly twice
as long as the coxa, the tibia not half as long as the femur. Tibia with four dorso-
apical teeth, and one ventral tooth. Tarsal segments in ratio 12:5 :5 :5 :7. Mid
leg slender, the subclavate tibia nearly as long as femur and trochanter combined;
carsal segments in ratio 20:10:9:8:10, long pubescent. Hind leg robust, the
coxa as long as the tibia, but much wider, the femur not quite twice as long.
Tibia (fig. 202) with two ventro-apical teeth, the antaxial one of which is biden-
tate. Tarsus pubescent, especially along the plantar edge; the segments in ratio
LOR OWE CES:
Gaster. Ovipositor distinctly projecting beyond the apex of the gaster.
Length, 2.6 mm. Colour of head and thorax dark brown; abdomen and extre-
mities lighter. Some females have the last three antennal segments, and the
distal segments of the legs, of an ivory-white colour, and are evidently not fully
mature. Nervures of the wings light brown, three dark striae radiate from the
stigma,
Remark.
C. iodotrichae is remarkable in having the apical segment in the male antenna
very long. The specimens are rather large, as in C. vechti, but they differ
distinctly in the relative proportions.
54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Ceratosolen notus (Baker), C. orientalis sp. n. and C. calopilinae sp. n.
Ceratosolen notus (Baker), and the new species C. orientalis and C. calopilinae
(described below) are very similar.
There are no distinct structural differences between C. orientalis and C. calopi-
linae, yet the two are recognizable by the male trophi, and by the relative propor-
tions of some body-parts, notably the female maxilla and antenna, and the male
hind leg.
C. orientalis is much like C. notus, but it differs in the relative proportions of
the hind leg in the male, of the female antenna, etc.
Examination of more material may prove the three forms to be only sub-
specifically distinct.
Ceratosolen notus (Baker) (figs. 204—217)
Blastophaga nota Baker, 1913, pp. 65—67, 69, 70—72, 81, 82, figs. 2, 3D, E, 4D [descr.
®, 6, ex Ficus nota (Blanco) Merrill, Los Bafios (Luzon, Philippine Is.), biological notes].
Ceratosolen notus: Williams, 1928, pp. 9—11, 14, PI. I figs. 2, 5, Pl. II fig. 16 [biological
notes].
Material.
Series 9, &, ex Ficus nota (Blanco) Merr., Los Bafios (Luzon, Philippine Is.),
Coll. Agric., leg. F. X. Wirrrams, 24.VI.1921; coll. HSPA, 9, & slide mounted;
Goll IIL, no NC MONO IO a,
Six 9, 8 &, ex Ficus nota (Blanco) Merr., Los Baños (Luzon, Philippine Is.),
leg. F. X. WILLIAMS, 26.11.1921; coll. HSPA; 2 9, 6 &, coll. USNM, 1 &
slide mounted.
Series!) 9, &, ex Ficus nota (Blanco) Merr., Los Baños (Luzon, Philippine
Is.), Coll. Agric., 9.VI.1921; leg. F. X. WILLIAMS, coll. HSPA.
Series 9, &, ex Ficus nota (Blanco) Merr. (several trees), Los Bafios (Luzon,
Philippine Is.), leg. F. X. WILLIAMS, 15/17.VI.1921; coll. HSPA.
Series 9, &, ex Ficus spec., Luzon (Philippine Is.), 20 min. SW of cou
leg. H. E. MILLIRON, 24.IX.1945; coll. BMH.
Series 9, 7 3, ex Ficus nota (Blanco) Merr., Manila (Luzon, Philippine Is.),
leg. D. T. FULLAWAY, 1.1.1921; coll. HSPA; 6.9, 2 &, coll. USNM, 1 2 slide
mounted.
Series @, 4, ex Ficus nota (Blanco) Merr. (“purple fruited”), Mt. Maquiling
(Luzon, Philippine Is.), 3000 ft. alt. leg. F. X. WırLıams, 25.1.1922; coll. HSPA.
One 4, ex Ficus nota (Blanco) Merr. (“purple 3”), Mt. Maquiling (Luzon,
Philippine Is.), at mud spring, leg. F. X. WirriaMms, 8.11.1922; coll. HSPA.
Series 9, 4, ex Ficus satterthwaitei2), Mt. Maquiling (Luzon, Philippine Is.),
leg. F. X. WILLIAMS, 21.IX.1921; coll. HSPA.
1) Except for three or four escapes, this is the total content of one fig: 759 9, 255 4,
no parasites or inquilines.
2) Ficus satterthwaitei Elmer = F. congesta Roxb.
J. T. WIEBES : Indo-Australian Ceratosolen 55
205
Figs. 204— 217, Ceratosolen notus, Luzon, 204, female fore tibia, 205—207, male, 205,
mandible, 206, detail of mid tibia, and tarsus, 207, apex of hind tibia, 208, female hind
tibia and metatarsus, 209-212, male, 209, hind leg, 210, fore tarsus, axial aspect, 211,
fore tibia and tarsus, 212, male, 213, female mandible, 214, female maxilla, 215—217, male,
215, cercus of tenth urite, and paramere, 216, maxilla (ventral aspect), and labium
(lateral aspect), 217, antenna. Figs. 209, 212, X 40; 204—208, 210, 211, 213, 214, 217,
X 115; 215, 216, X 275
IDES CRP Un Os,
As the original description by BAKER is very short, I give a redescription of
C. notus. The described specimens, taken at the type locality of C. notus (College
of Agriculture, Los Bafios, WILLIAMS, 24.VI.1921), were sent to me dry, and I
relaxed them in diluted alcohol.
Male. Head (fig. 212) more than twice (9 : 4) as long as wide anteriorly, the
maximum width two-thirds of the length. Eyes absent. Epistomal margin wide,
with four hairs. Pubescence long. Antennal grooves open, rather wide behind.
Antenna (fig. 217) five-segmented: the scape, nearly twice as long as wide, one
and a half times as long as the pedicel (2 : 1): the first flagellar segment short,
the second (5:4) three quarters the length of the pedicel, the apical segment
(4:1) two and a half times as long as the second. Maxilla (fig. 216) expanded
laterally, with two lateral hairs; labium with one subapical hair. Mandible,
fig. 205.
56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Thorax, fig. 212. Pronotum slightly longer than wide posteriorly, and not quite |
twice as long as wide anteriorly. Mesonotum (8 : 5) much wider than long; meta- |
notum narrower, incompletely separated from the subquadrate propodeum. Fore
leg (figs. 210—211): the tibia not quite half as long as the femur [7 : 16},
apically with three dorsal and three ventral teeth. Tarsus bimerous, the distal
segment slightly longer than the proximal one. Mid leg rather robust; coxa and |
femur subequal in length; the tibia distinctly longer [4:3], with four apical |
teeth (fig. 206). Tarsus coiled in nearly all specimens, the segments approximately
in ratio 5:3 :3:4:10. Hind leg (fig. 209) pubescent, the coxa three quarters
the length of the femur, the tibia slightly longer than the coxa. Tibial armature |
(fig. 207) consisting of a bidentate antaxial process, a small tooth at the ventral
edge, and a motile axial spur. Tarsus dilated, the pubescence slightly longer than |
one third of the width of the segments. Relative proportions of the segments |
rather variable, even in specimens from the same sample. First segment always |
approximately half as long as the tibia, second and third segments subequal, ap- |
proximately half as long as the first; the fourth distinctly longer than the third;
the fifth segment is slightly longer (as in fig. 209) or much longer and wider
(as in BAKER's fig. 3D) than the fourth.
Gaster. Cerci of the tenth urite with three claws, parameres narrow (fig. 215).
Length, 1.6 mm. Colour yellowish brown.
Female. Scape of the antenna large, nearly four times as long as the pedicel,
which bears some thirty slender spines at the axial surface. Appendage of the
third segment robust, the fourth segment small. Fifth to eighth segments
gradually increasing in length, the eighth distinctly longer than the fifth [5 : 4];
the ninth segment as long as the fifth, the tenth slightly longer, the eleventh,
forming a loose club with the tenth, distinctly shorter [3 : 4]. Funicular segments
with two irregular rows of short, wide sensilla. Mandible, and its appendage, with
five ventral ridges (fig. 213). Maxilla (fig. 214) with a subapical hair, and a
bacilliform process, which is two-fifths the length of the maxilla.
Thorax pubescent; two hairs occur above, and a group of about ten hairs below
the propodeal spiracle. Fore wing (5 :2), 1.8 mm long. Submarginal, marginal,
stigmal, and postmarginal veins in ratio 28 : 10 : 9 : 24; submarginal and stigmal
veins with four pustules each. Hind wing (5:1), 1.1 mm long. Tibia of the
fore leg (fig. 204) less than half as long as the femur {5 : 13], and more than
half as long as the coxa [5 : 8}. Apical armature consisting of a dorsal comb of
four teeth, and one ventral tooth. Tarsal segments in ratio 20:10:9:8:15.
Mid leg long pubescent, the tibia shorter than femur and trochanter combined
[10 : 11}; tarsal segments in ratio 18 : 8 :9 : 8 : 14. Coxa of hind leg (fig. 208)
shorter than the femur [14:17], the tibia two-thirds the length of the femur.
Antaxial apical tooth bidentate, the axial tooth simple, strongly curved at apex.
Tarsus heavily pubescent and spinose, the segments in ratio 20:7:7:6:9.
Gaster. Projecting part of the ovipositor one sixth the length of the gaster.
Length, 1.8 mm. Colour brown, head darker, legs yellow-brown. Nervures of
the wings light brown.
J. T. Wiegers: Indo-Australian Ceratosolen 57
Remark.
There are some minor differences between the specimens described above and
the description by BAKER, notably in the male genitalia (described by BAKER
as being “without armed claspers”, but the cerci are not distinctly visible) and in
the relative proportions of the female hind leg (tibia half as long as the femur,
according to BAKER, two-thirds in length of the femur in the specimens
measured by me, but probably BAKER measured the part of the tibia visible in
situ, and not the whole segment).
| I fail to find any differences between the specimens from Ficus nota and those
from F. congesta. For a discussion of the host records, see p. 97.
Ceratosolen orientalis sp. n. (figs. 218—230)
Material.
Series 9, 4, ex Ficus subcongesta Corner (det. E. J. H. CORNER), Keravat
(New Britain), leg. E. J. H. CORNER, 14.X.1960, NGF no. 13783; coll. no.
540; & (holotype), slide 540a, 2 (allotype), 540c, &, 9 (paratypes), 540b, d.
? Sixteen 19, 8 4, ex Ficus subcongesta Corner var. symmetrica Corner (det.
E. J. H. CORNER), Crown Prince Mts. (Bougainville I.), leg. E. J. H. CORNER,
ING Elino #5 755 coll now 50s nd. slidess5 Oan 2, 550b.
Description.
Male. Head (fig. 222) slightly longer than twice the anterior width, maximum
width two-thirds of the length. Pubescence short and sparse. Eyes small. An-
tennal grooves open, not very wide behind. Antenna (fig. 220): the scape
(5 :3) is two and a half times as long as the pedicel (1 :1), which is twice as
long as the first flagellar segment (1 : 2). The following segment (8 : 7) as long
as the pedicel, and not quite half as long as the apical one (3:1). Maxilla (fig.
230) with two lateral hairs; labium without long hairs. Mandible, fig. 224.
Thorax, fig. 222. Pronotum nearly twice as long as wide anteriorly; the
posterior width slightly shorter than the length. Mesonotum transverse (14 : 9);
metanotum incompletely separated from the propodeum, which is nearly twice as
wide as long. Femur of fore leg (fig. 225) twice as long as the tibia (apical teeth
included); the tibia with four dorsal and three ventro-apical teeth. Tarsus
bimerous, segments in ratio 5 : 6. Coxa of the mid leg as long as the femur, the
tibia as long as femur and trochanter combined, subclavate, with four apical teeth.
Hind leg (figs. 223, 227): coxa and femur subequal in length, the tibia little
shorter. Tibia with a bidentate process at the antaxial apical edge. Tarsus dilated,
segments approximately in ratio 6 : 4 : 4 :3 : 5. Pubescence longer than half the
width of the segments.
Gaster. Cerci of the tenth urite with three or four claws.
Length, 1.5 mm. Colour uniformly yellowish brown.
Female. Head shorter than wide across the compound eyes [15 : 16}. Eye longer
than the cheek [11 :9}. Pubescence short and sparse. Antenna (fig. 219): scape
not very wide, and nearly thrice as long as the pedicel, which bears forty-five
axial spines. Third segment not very long, the fourth small; the fifth twice, the
58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
in
it
Figs. 218—230, Ceratosolen orientalis, New Britain (228, specimen from Bougainville I.),
218, female mandible, 219, female antenna, 220, male antenna, 221, female fore leg,
222—225, male, 222, male, 223, hind leg, 224, mandible, 225, detail of fore leg, 226,
female maxilla, 227, detail of male hind leg, 228, do., specimen from Bougainville I., 229,
detail of female hind leg, 230, male labium and maxilla. Figs. 222, 223, X 50; 221, 225,
229, X 115; 218—220, 224, 226—228, X 155; 230, X 365
J. T. Wiegers: Indo-Australian Ceratosolen 59
sixth and seventh approximately thrice as long as the fourth. Eighth to eleventh
segments slightly shorter than the seventh, and gradually diminishing in length.
The subapical and apical segments shaped so as to form a loose club. Fifth segment
with eight oblong sensilla, sixth to tenth segments with two irregular rows of
sensilla (eleven to fourteen per segment), the apical segment with six oblong, and
some circular sensilla. Maxilla (fig. 226) with one subapical hair, and a long
bacilliform process. Mandible (fig. 218), and its appendage, with five ventral
ridges.
Thorax. Pronotum with long hairs. Scutum longer than wide posteriorly [4 : 3},
the posterior width slightly longer than half the maximum width. Lateral margins
with six hairs. Scutellum one and a half times as long as wide anteriorly, and
slightly longer than wide posteriorly; with rows of six or seven hairs along the
lateral margins, and approximately ten hairs on the disk. Metanotum rather long,
with six hairs on each side of the mid line. Propodeum with two hairs above,
and four beneath the large spiracle. Fore wing (7 : 3), 1.7 mm long. Submarginal,
marginal, stigmal, and postmarginal veins approximately in ratio 3 : 1 : 1 : 2; sub-
marginal vein with three pustules, stigmal vein with four. Hind wing (5:1),
0.9 mm long. Femur of the fore leg (fig. 221) longer than the coxa [3 : 2}, and
thrice as long as the tibia. Tibia with a dorso-apical row of four teeth, and one
ventral tooth. Tarsal segments in ratio 20 :4:4:5 :10. Tibia of the mid leg as
long as femur and trochanter combined, with long hairs. First tarsal segment as
long as the following two combined, second to fifth segments gradually, but
slightly, increasing in length. Femur of the hind leg (fig. 229) slightly longer
than the coxa, the tibia two-thirds the length of the femur, with two apical teeth,
the antaxial one of which is bidentate. Tarsus rather pubescent, especially along the
plantar edge; segments in ratio 12:6:5:5:6.
Gaster. Ovipositor slightly projecting beyond the apex of the gaster.
Length, 1.9 mm. Colour brown, ventral surface and extremities lighter. Ner-
vures of the wings brown, stigma with dark radiating striae.
Remark.
The specimens from Bougainville I. (coll. no. 550) differ from those of the
typical sample. The females have the third antennal segment more slender, with
longer and more slender, hyaline processes, and the pedicel bears less (approximate-
ly twenty-five) teeth at the axial surface. In the male, the first flagellar segment
of the antenna is only one third the length of the second. The hind leg is shorter
than in the New Britain specimens, and the metatarsus is less slender. The tibia
bears a prominent tooth at the axial edge (fig. 228). As the specimens are few,
and not in a very good condition, I do not name the form from Bougainville I.,
which should perhaps be regarded as a distinct species or subspecies.
Ceratosolen calopilinae sp. n. (figs. 231—242)
Material.
Series 9, 4, ex Ficus calopilina Diels (det. E. J. H. CORNER), Wau (Terr.
New Guinea), leg. E. J. H. CORNER, 13.IX.1960; coll. no. 538; & (holotype),
slide 538a, © (allotype), 538c, 4, 9 (paratypes), 538b, d, e.
60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Dieser ip’ Bilon. |
Male. Head (fig. 237) little more than twice as long as wide anteriorly, and |
nearly twice as long (8 : 5) as the maximum width. Eyes absent. Antennal grooves
open, not very wide behind. Antenna (fig. 238) slender; the scape (2 : 1) nearly
twice as long as the pedicel (3 : 2), the first flagellar segment short, the second
(7:6) distinctly shorter than half the length of the apical segment (4:1). |
Mandible, fig. 236. Labium and maxillae, fig. 241; maxilla with one lateral hair. |
Thorax, fig. 237. Pronotum twice as long as wide anteriorly, and one and a half
times as long as wide posteriorly. Mesonotum with a straight posterior margin, |
tapering anteriorly; the length more than half the maximum width. Metanotum
which is approximately one and a half times as long as wide. Fore leg (fig. 240): |
the tibia nearly half as long as the femur, with three dorsal and three ventro-
apical teeth. Tarsus bimerous, distal segment distinctly longer than the proximal |
one, with long hairs. Coxa and femur of mid leg subequal, approximately as long |
as the tibia. Tibia subclavate, with long hairs along the dorsal margin and on the.
axial surface. The apical edge dorsally produced into a stout tooth, ventrally with
four smaller teeth. Tarsal segments in ratio 8:4:4:3:11. Coxa of the hind
leg (fig. 232) little shorter than the femur. Tibia with one apical antaxial tooth,
pubescent along the dorsal edge. Tarsus dilated, with pubescence approximately
half as long as the width of the segments. There is some variation in the dimen-
sions of the segments, but the relative lengths are rather constant: 11 : 8:7: 6:10.
Gaster. Aedeagus slightly dilated at mid length. Cercus of the tenth urite with
three claws, paramere with parallel edges (fig. 239).
Length, 1.3 mm. Colour uniformly yellow-brown.
Female. Head slightly shorter than wide across the compound eyes. Longitudinal
diameter of the eye longer than the cheek [8:7]. Pubescence short. Antenna
(fig. 231): scape more than twice as long as the pedicel, which bears approxim-
ately twenty axial spines. Third segment not very long, the fourth small, the fifth
shorter than the sixth {5 : 6}, the seventh longer than the sixth {7 : 6}. Eighth to
tenth segments gradually diminishing in length, the tenth as long as the sixth.
Eleventh and tenth segments shaped so as to form a club, but not fused. Fifth seg-
ment with ten sensilla, sixth to tenth segments with thirteen or fourteen sensilla
in two irregular rows; the apical segment with six oblong and two circular sensilla.
Mandible (fig. 235), and its appendage, with six ventral ridges. Maxilla (fig.
234): bacilliform process not quite half as long as the maxilla [4:9}. Labium
without long hairs.
Thorax. Pronotum wide, with long hairs. Scutum one and a half times as long
as wide posteriorly, and nearly twice as wide anteriorly; with rows of four hairs
along the lateral margins. Scutellum one and a half times as long as wide, with
rows of five hairs along the lateral margins, and approximately ten hairs on the.
disk. Metanotum short, with nine hairs on each side of the mid line. Propodeum |
with three hairs above, and six beneath the propodeal spiracle. Fore wing (7 :3),
1.6 mm long. Submarginal, marginal, stigmal, and postmarginal veins in ratio |
75 :17 : 24 : 44. Submarginal vein with three pustules, stigmal vein with five.
J. T. WieBEs: Indo-Australian Ceratosolen 61
Figs. 231— 242, Ceratosolen calopilinae, New Guinea, 231, female antenna, 232, male hind
leg, 233— 235, female, 233, detail of fore leg, 234, maxilla, 235, mandible, 236— 241, male,
236, mandible, 237, male, 238, antenna, ventral aspect, 239, cercus of tenth urite, and
paramere, 240, detail of fore leg, 241, labium and maxillae, 242, detail of female hind leg.
Figs. 232, 237, X 50; 231, 233, 240, 242, X 115; 234—236, 238, X 155; 239, 241, X 365
Hind wing (4:1), 0.9 mm long. Coxa of the fore leg (fig. 233) more than half
as long as the femur, tibia distinctly smaller than the coxa. Tibia with four dorso-
apical teeth, and one ventral tooth. Tarsal segments in ratio 13 :4:4:5 :6. Mid
leg of the usual slender build, the clavate tibia slightly longer than the femur;
tarsal segments in ratio 15 : 7 :7 : 5 :9. Coxa of the hind leg (fig. 242) shorter
than the femur [4:5]. Tibia nearly three quarters the length of the femur, with
two ventro-apical teeth, the antaxial one of which is bidentate. Tarsus pubescent,
segments in ratio 25:10:8:8:11.
Gaster. Ovipositor short, but distinctly projecting beyond the apex of the
gaster.
62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Length, 1.8 mm. Colour dark brown, ventral surface and extremities lighter. |
Nervures of wings light brown.
Remark.
C. calopilinae is very similar to C. orientalis, but it differs in the length of the
bacilliform process of the female maxilla, the chaetotaxy of the male maxilla, and
in the proportions of the male hind leg.
Ceratosolen corneri sp. n. (figs. 243—254)
Material.
Series 9, 4, ex Ficus endothrix1), Mt. Maquiling (Luzon, Philippine Is.), at !
first creek crossing, leg. F. X. WILLIAMS, 27.VI.1928, no. 3131; coll. HSPA, & |
(holotype), 9 (allotype), slide mounted; coll. ML, no. 682: 10 9, 10 4.
Fragments of several &, and 1 9, ex Ficus botryocarpa Miq. var. subalbido-
ramea (Elmer) Corner (det. E. J. H. CORNER), Mindoro (Philippine Is.), leg.
E. D. MERRILL, no. 1813; coll. no. 405; & (paratypes), slide 405a, 9 (paratype),
405b.
Series 9, &, ex Ficus barnesii2), Mt. Maquiling (Luzon, Philippine Is.), leg.
F. X. WILLIAMS, 28.VI.1928, no. 31313), coll. HSPA.
Description.
Male. Head (fig. 246) more than twice as long as wide anteriorly. Epistomal
margin (fig. 247) with four hairs. Eyes very small. Antennal grooves open,
narrow behind. Antenna (fig. 248) five-segmented, the scape (5 :2) four-thirds
the length of the pedicel (15 : 7). Pedicel six times as long as the first flagellar
segment (5 : 13), the second segment (25 : 14) half as long as the apical segment
(4:1), and five times as long as the first. Maxillae (fig. 252) for a great part
concealed under the hypostomal margin; labium atrophied. Mandible, fig. 251.
Thorax, fig. 246. Pronotum more than twice as long as wide anteriorly, its
length one and a half times the maximum width. Mesonotum wider than long
(7:5). Metanotum incompletely separated from the propodeum, not nearly as
wide as the mesonotum [11:14]. Propodeum shorter than wide {3 : 4]. Fore
leg (fig. 243): the tibia not quite half as long as the femur, with four dorsal and
two ventro-apical teeth. Tarsal segments subequal in length. Coxa of the mid
leg longer than the femur; the tibia longer than the femur, but not as long as the
coxa. Tibia with four sharp apical teeth. Tarsal segments in ratio 22 :6:5:4:14.
Hind leg (fig. 253) very long, all segments with thick pubescence of small hairs.
Coxa and femur very long (each about as long as the pronotum), and dilated.
Tibia more than half as long as the femur, without any distinct apical armature.
Tarsal segments in ratio 15 :8 :7 : 6 : 7. Metatarsus distinctly produced dorsally.
Gaster. Cerci of the tenth urite (fig. 249) with three claws.
Length, 1.5 mm. Colour uniformly yellow-brown.
1) Ficus endothrix Warb. = F. botryocarpa Miq.
2) Ficus barnesii Merr. = F. botryocarpa Miq.
3) ?, same tree as the first sample (no. 682) ?
J. T. WIEBES : Indo-Australian Ceratosolen
248
251
Figs. 243—254, Ceratosolen corneri, Luzon (except 246, 247, 251, 254, specimen from
Mindoro), 243, detail of male fore leg, 244, detail of female hind leg, 245, female mandible,
246—249, male, 246, male, 247, epistomal margin, 248, antenna, 249, cercus of tenth urite,
and paramere, 250, female fore leg, 251—254, male, 251, mandible, 252, maxillae and
hypostomal margin, 253, detail of hind leg, 254, hind leg (specimen from Mindoro). Figs.
246, X 50; 253, 254, X 80; 243, 244, 247, 248, 250, X 155; 245, 251, X 215; 249,
PSD. YK BOSD
64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Female. Antenna consisting of eleven segments, the last two of which are
joined into a loose club. Scape twice as long as the pedicel, appendage of the
third segment robust, the fourth segment small. Length of the fifth segment
four-fifths of that of the sixth, two-thirds of that of the seventh, eighth and tenth
segments; the ninth as long as the sixth segment. Eleventh segment half as long
as the tenth. Funicular segments with one or two rows of long sensilla. Maxilla
with one subapical hair, without a bacilliform process. Mandible (fig. 245) with
five ventral ridges, its appendage with five large ridges, and smaller ridges in be-
tween the larger ones.
Thorax. A few hairs occur along the lateral edges of the scutum, five hairs
along the edges of the scutellum. Metanotum with about ten hairs on each side
of the mid line. Propodeum with two hairs above, and five beneath the spiracle.
Fore wing (5 : 2), 1.4 mm long. Submarginal, marginal, stigmal, and postmarginal
veins in ratio 12 : 4 :3 : 6; the submarginal and stigmal veins with three pustules
each. Hind wing (5 : 1), 0.8 mm long. Fore leg (fig. 250): the coxa smaller than
the femur [11 : 14}, and approximately twice as long as the tibia. Tibial armature
consisting of four dorsal teeth. Tarsal segments in ratio 8 :3:3:4:5. Mid leg
slender, the tibia as long as femur and trochanter combined, tarsal segments in
ratio 25 :9:9:7:10. Coxa of hind leg (fig. 244) six-sevenths the length of the
femur, tibia slightly longer than half the length of the femur. Tibial armature
consisting of a bidentate antaxial, and a long, curved axial tooth. Tarsus fimbriated
at the plantar edge, segments in ratio 8:3:2:2:3.
Gaster. Projecting part of the ovipositor not quite one quarter the length of the
gaster.
Length, 1.4—1.6 mm. Colour brown, ventral surface and extremities lighter.
Nervures of the wings light brown.
Remark.
Although the female of this species is in no way aberrant from the normal
type, the male is very remarkable. Its very long hind legs distinguish it at once
from its congeners; in fact, in none of the other fig wasps have I seen the like.
There are some differences between the samples from Luzon and Mindoro. The
specimens from the last mentioned locality are badly damaged, and better series
should be studied to be certain of the differences. The males have the tarsus of
the hind leg slightly more robust, and the metatarsus without the dorsal protuber-
ance (fig. 254).
I name this species in honour of Mr. E. J. H. Corner FRS. Lecturer in
Botany, University of Cambridge.
Ceratosolen solmsi (Mayr) and C. marchali Mayr (figs. 255—260, 262—265;
map, fig. 261; Table I)
Two forms of Ceratosolen were described that are known to live in Ficus
hispida Linn. f., viz., C. solmsi (Mayr) and C. marchali Mayr. According to
GRANDI (1928c, pp. 174—175), the species differ in the following characters:
9. Eleventh antennal segment longer than the tenth in C. solmsz; shorter in
C. marchali.
J. T. WIEBES : Indo-Australian Ceratosolen 65
Bacilliform process of the maxilla one quarter the length of the maxilla in C.
solmsi; one half as long as the maxilla in C. marchali.
? Mandibular process with six or seven ridges in C. solmsi; with nine ridges
in C. marchali.
Fore wing hyaline in C. 50/7257; darker, with striae radiating from the stigmal
vein, in C. marchalt.
&. Antennal groove narrow behind in C. solmsz; wide in C. marchali.
Fourth antennal segment slightly longer than the third in C. solmsi; distinctly
longer in C. marchalı.
According to GRANDI (1926, p. 358), the male hind tibia is thrice as long as
the first tarsal segment in C. solmsi from Java; and twice as long in C. marchali
from Vietnam.
The forms were known from Java and S. Vietnam (C. solmsi) and from
N. Vietnam and India (C. marchali). Moreover, GRANDI (1928c, p. 176) found
one slide in Mayr’s collection, labelled “Tonkin”, and containing C. solmsi.
I have seen material from Ceylon, India, Malaya (Selangor and Langkawi Is.),
Hongkong, Java, and Queensland (distribution, see map, fig. 261). The data
on some of the above mentioned differential characters are arranged in Table 1
(cf. figs. 255—269).
The male mouthparts are identical in the two forms (fig. 264). The genitalia
bear cerci (fig. 265), which are clearly visible in the specimens from the Asian
continent. In the specimens from Java and Langkawi, however, they are hyaline,
and very difficult to be observed: I cannot even find them in all of these spec-
imens, and in none of the sample from Queensland (but this is badly preserved).
In general, the specimens from India, Ceylon, and Hongkong are slightly smaller
and less slender than those from Malaya, Java, and Queensland.
It seems best to treat C. solmsi and C. marchali as subspecies. The specimens
from Java are distinctly different from those collected in Ceylon, India, and
China, but the characters seem to intergrade on the Malayan peninsula. The
Australian specimens!) are intermediate in some aspects, but they resemble C.
marchali more closely than they do C. solmsz.
Ceratosolen solmsi solmsi (Mayr)
Blastophaga (Ceratosolen) Solmsi Mayr, 1885, pp. 154, 161, 163, 168—169, Pl. IX figs.
4—5 [key 9, &, descr. 9, &, ex Ficus (Cystogyne) canescens Kurz?) (det. Solms), Bogor
(Java), Bot. Gdn., leg. H. Solms-Laubach; type: @ ].
Ceratosolen Solmsi: Mayr, 1906, p. 155 [rectification of original descr., ex Covellia didyma
Miq.), Bogor (Java), Bot. Gdn., leg. C. Aurivillius, no. 2}; Grandi, 1928c, pp. 173—176
{type specimens of Blastophaga Solmsi Mayr studied, = C ? Marchali: Grandi; differences
from C. marchali Mayr}.
Ceratosolen ? Marchali: Grandi, 1926, pp. 357—358 [2, &, ex Ficus hispida L. (det.
J. Beumée; Herb. Bog. no. E. J. 2173), Djakarta (Java), leg. E. Jacobson, IV.1924]; Grandi,
1927b, pp. 174—178, figs. III—IV [descr. 9, 4, ex Ficus spec., Saigon (S. Vietnam), leg
F. Silvestri, 1924].
1) Series 9, 4, ex Ficus hispida Linn.f., Hambledon (Queensland), leg. C. E.
PEMBERTON, 9.X1.1921; coll. HSPA; coll. ML, no. 590: 20 ®, 10 4.
2) For a discussion of the host records, see p. 97.
66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
255 256
Figs. 255—260, Ceratosolen s. solmsi (255, 256) and C. s. marchali (257—260), antennal
groove of male, specimens from: 255, Ceylon, 256, 257, Selangor, 258, Langkawi I., 259,
Java, 260, Queensland, 261, distribution on the Asian continent, and in the Philippines and
Indonesia, of: C. s. solmsi (black triangles), C. s. marchali (open triangles) — the numbers
refer to the figures above —, C. bisulcatus (black dots), C. jucundus (open circles), C.
constrictus (black squares), and C. hewitti (open squares), 262—264, C. s. solmsi, Java,
male, 262, detail of right antenna, with the measurements of third and fourth segments,
used in Table I, 263, detail of hind leg, with the measurements of tibia and metatarsus, used
in Table I, 264, labium and maxillae, 265, C. s. marchali, Selangor, male, cercus of tenth
urite. Figs. 255—260, 263, X 105; 265, X 140; 264, X 190; 262, X 235
Material.
Series 9, &, ex Ficus hispida Linn. f., Bogor (Java), Bot. Gdn, VIII.1954;
coll MO, 9.10, 112 (20S re 47 Gy alll ion Ons die), 125 8, slice 2a,
One 2,1 &, Java, no. 186; coll. USNM.
Fragments of 5 9, series &, ex Ficus hispida Linn. f.1) (det. E. J. H. Cor-
NER), Langkawi Is. (Malaya), 1941, Sing. F. no. 37889; coll. no. 412; 9, slide
41 2a.
1) “This was a rather unusual form of F. hispida with completely geocarpic figs (borne
on slender twigs rooting in the ground)” (CORNER, 1956, in litt.).
67
Indo-Australian Ceratosolen
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68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Ceratosolen solmsi marchali Mayr
Ceratosolen Marchali Mayr, 1906, pp. 155—156 [descr. 9, 4, ex Ficus spec, Tonkin |
(N. Vietnam) leg. P. Marchal, no. 20a}; Grandi, 1928c, pp. 173—175 [type specimens of
C. Marchali Mayr studied, = C. Berlandi Grandi; differences from C. solmsi (Mayr) }:
Joseph, 1953c, p. 282 [ 2, 6, ex Ficus hispida L., Kottayam (Travancore, India), leg. K. J.
Joseph, 26.VI.1951].
Ceratosolen Berlandi Grandi, 1928a, pp. 74—79, fig. II [descr. 9, &, ex Ficus hispida
L., Chambaganour (Pulneys, India)!), 5000 ft. alt. leg. E. Gombert, 14.IV.1914].
Material.
Series 9, &, India, ex coll. JosEPH; coll. ML, no. 564.
Ten 3,5 9, Calcutta (India), leg. ROTHNEY, VIII.1885; coll. OUM.
Series 9, 4, ex Ficus hispida Linn. f., Hongkong (China), leg. GREEN, is-
sued at Honolulu, 29. VIII—1.IX.1921; coll. HSPA; coll. ML, no. 594:1 9,1 4.
Series 9, 4, ex Ficus hispida Linn. f., Hongkong (China), leg. D. T. FULLA-
WAY, 16.1.1921; coll. HSPA.
Series 9, &, ex Ficus hispida Linn. f. (det. E. J. H. CORNER), Sungei Gombah
(Selangor, Malaya), leg. E. J. H. CORNER, 4.X.1961; coll. no. 618; 9, slide 618a,
4, 6185, &
Ten 9, 2 &, ex Ficus hispida Linn. f., Peradeniya (Ceylon), Bot. Gdn., leg
F. KEISER, 25.11.1954: coll. NMB: coll. MIL, no. 378; 1 ©.
Three 9, series &, ex Ficus hispida Linn. f., Peradeniya (Ceylon), Bot. Gdn.,
leg. F. KEISER; coll. NMB; coll. ML, no. 382: 3 &.
Ceratosolen brongersmai sp. n. (figs. 266—277)
Material.
Series 9, 4, ex Ficus treubii King (det. E. J. H. CORNER), Sungei Mesilau
(N. Borneo), 5500 ft. alt leg. E. J. H. Corner, VIII.1961; coll. no. 604; 4
(holotype), slide 604a, @ (allotype), 604c, 4, ® (paratypes), 604b, d.
Series 9, &, ex Ficus treubii King (det. E. J. H. CORNER), Sungei Bembangan
(N. Borneo), 5500 ft. alt., leg. E. J. H. Corner, VIII.1961, RSNB no. 1891;
Gull, sos GV? @, chide 0123. @ 8, Go
Desert DIEKE
Male. Maximum width of the head (fig. 269) three quarters of the length,
the head slightly narrower anteriorly and posteriorly. Pubescence short and sparse.
Eyes absent. Antennal grooves open, wide behind. Antenna (fig. 270) five-
segmented: the scape (7 : 4) one and a half times as long as the pedicel (2:1);
the first flagellar segment annuliform, the second (1 : 2) less than one fifth the
length of the third (3 : 1). Labium and maxillae (fig. 277) large, the labium bilo-
bate, the lobes not equal; maxilla with a lateral expansion bearing one long hair.
Mandible, fig. 266.
Thorax, fig. 269. Pronotum nearly twice as long as wide anteriorly, the
posterior width one and a half times the anterior width. Maximum width of the
1) ? = Shembaganur, Palni Hills, S. India?
J. T. WieBEs: Indo-Australian Ceratosolen 69
mesonotum, across the lateral angles, distinctly longer than the length [13 : 10}.
Metanotum narrower than the mesonotum, incompletely separated from the pro-
podeum. Propodeum long and rather narrow, with large peritremata. Metasternum
glabrous. Femur of the fore leg (fig. 272) twice as long as the tibia. Tibia apically
with four dorsal teeth (three large and one smaller), and three ventral teeth.
Tarsus bimerous, segments in ratio 5 : 6. Mid leg slender, the coxa nearly as long
as the femur, the tibia distinctly longer [3 : 2}. Tibia with the ventro-apical edge
produced into two teeth. Tarsus pentamerous, the segments in ratio 7:2:2:2:
7. Hind leg, fig. 267. Coxa large, with a hyaline border along the dorsal margin as
in the species with dilated hind feet, pubescent axially, glabrous antaxially. Femur
with a hyaline part along the distal ventral border; antaxial surface with small
hairs, axially with many long stout hairs. Tibia two-thirds the length of the femur;
ventral apex with two articulating teeth: the axial one curved, the antaxial one
deeply bidentate. Tarsus with small protuberances along the ventral margin; the
five segments in ratio 16:6:5:5:14.
Gaster. Cerci of the tenth urite with three claws, parameres straight and narrow
(fig. 273).
Length, 1.7 mm. Colour uniformly olive-brown, the pubescent parts of the hind
legs darker.
Female. Head as long as wide across the eyes. Longitudinal diameter of the eye
longer than the cheek [7 : 5}. Pubescence long. Antenna (fig. 271): scape not
very long, but wide; the pedicel not quite half as long as the scape, with approxi-
mately thirty-five spines at the axial surface. Appendix of the third segment long
and slender; the fourth segment small. Fifth segment wide, with twelve long
sensilla. Sixth segment longer than the following [8 : 7}, seventh to ninth seg-
ments subequal, and longer than the tenth [7 : 6}, which is a little longer than
the apical segment. Sixth to tenth segments with two irregular rows of ten long
sensilla each; the eleventh segment with ten long and two circular sensilla. Labium
and maxillae (fig. 276): labium with two hairs, maxilla with one apical hair;
maxilla moreover with a bacilliform process, which is nearly half as long as the
maxilla. The process bears two hairs. Mandible (fig. 275) with four large and
four smaller ventral ridges, the appendage with six or seven ventral ridges.
Thorax. Pronotum not very large, pubescent. Scutum nearly as long as its maxi-
mum width, a little narrower behind; with approximately twenty long hairs. Scu-
tellum one and a half times as long as wide, with eight hairs along each lateral
edge. Metanotum with a row of nine or ten hairs on each side of the mid line.
Propodeum large, with two hairs above, and fifteen beneath the peritremata. Fore
wing (2:1), 2.0 mm long. Submarginal, marginal, stigmal, and postmarginal
veins in ratio 7:3:3:4; the stigma with four pustules, the submarginal vein
with three. Hind wing (4:1), 1.2 mm long. Fore leg (fig. 274): the coxa two-
thirds the length of the femur, which is more than twice as long as the tibia.
Apical tibial comb consisting of four sharp teeth. Tarsus pentamerous, the seg-
ments in ratio 18:8:7:7:8. Mid leg slender, with long hairs. Tibia nearly as
long as femur and trochanter combined, with a long, slightly curved ventral spur.
Tarsal segments in ratio 17 : 8:8: 8:11. Coxa of the hind leg (fig. 268) large;
70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Figs. 266—277, Ceratosolen brongersmai, Borneo, 266, male mandible, 267, male hind leg,
268, detail of female hind leg, 269, male, 270, male antenna, 271, female antenna, 272,
detail of male fore leg, 273, male, cercus of tenth urite, and paramere, 274— 276, female,
274, fore tibia, 275, mandible, 276, labium (lateral aspect), and maxilla, 277, male labium
and maxillae. Figs. 269, X 40; 272, X 90; 267, 268, 271, 274276, X 115; 266, 270)
x 1603 278; Zits 27/5
the femur long but not very wide, with long hairs; the tibia as long as the coxa, and
nearly two-thirds the length of the femur. Tibia with two ventral teeth: the axial
one rather long and curved, the antaxial one bidentate. Tarsus pubescent, especially
along the plantar edge; segments in ratio 15 :6:6:4:8.
Gaster. Projecting part of the ovipositor about one tenth the length of the
gaster.
Length, 2.0 mm. Colour brown, the extremities and ventral surface lighter.
Nervures of the wings brown, light brown striae radiate from the stigma.
J. T. WIEBES : Indo-Australian Ceratosolen zijl
Remark.
In some aspects, this species is close to C. humatus and C. albulus (see below),
notably in the shape of the male trophi. The female, however, has a different tibial
armature in the fore leg, and the male hind tarsi are not dilated. The other con-
nection is that with C. solmsi, which, however, has different male mouthparts. For
further remarks on the classification of C. brongersmai and C. solmsi, see p. 87.
I name this remarkable species in honour of Dr. L. D. BRONGERSMA, Director
of the Rijksmuseum van Natuurlijke Historie, Leiden.
Ceratosolen pilipes sp. n. (figs. 278—285; 287—292)
Material.
Series 2, &, ex Ficus cereicarpa Corner (det. E. J. H. CORNER), Mt. Kinabalu
East (N. Borneo), 2000 ft. alt. leg. E. J. H. CORNER, VI.1961; coll. no. 636; 4
(holotype), slide 636a, 2 (allotype), 636e, 4, ® (paratypes), 636b, c, d, f.
Diese epi tom:
Male. Head (fig. 279) nearly twice as long as wide anteriorly; the maximum
width two-thirds of the length. Dorsal surface, especially along each side of the
antennal grooves, with rather long hairs. Eyes present. Antennal grooves open,
wide behind. Antenna (fig. 281) five-segmented; the scape (17 : 10) nearly twice
as long as the pedicel (9 : 5); the first flagellar segment annuliform, the second
(1 : 1, longer antaxially than axially) one third the length of the apical segment
(9:4). Labium and maxillae, fig. 288. Labium with two subapical hairs, and
sometimes with a pair of basal hairs, which, however, more often than not are
absent. Maxilla expanded laterally, with two lateral hairs. Mandible, fig. 278.
Thorax, fig. 279. Pronotum not quite twice as long as wide anteriorly, and
nearly as long as wide posteriorly. Mesonotum large, little wider than long. Meta-
notum incompletely separated from the propodeum; propodeum long and rather
narrow, with large peritremata. The metasternum, and to a lesser extent also the
mesosternum, with a rather long and heavy pubescence (fig. 289). Fore leg (fig.
292): the coxa large, the femur twice as long as the tibia including the large dorsal
teeth. Tibia with a comb of four closely joined large teeth at the dorsal edge, and
three ventral teeth. Tarsus bimerous, but with an indication of a trimerous con-
dition; the two free segments subequal in length. Coxa of the mid leg little shorter
than the femur; the tibia as long as femur and trochanter combined, with heavy
pubescence apically, the ventral edge produced into two teeth. Tarsus pentamerous;
the segments in ratio 23:7:9:8:23. Hind leg (fig. 287) with heavy pubes-
cence on all segments. Coxa wide, but shorter than the femur. Tibia two-thirds the
length of the femur; the dorso-apical edge produced; ventrally with a bidentate
tooth. Tarsal segments in ratio 12:6:6:5 :9, the length of the pubescence ap-
proximately two-thirds the width of the segments.
Gaster partly sclerotized. Cerci of the tenth urite (fig. 284) with five or six
short claws, the parameres curved axially.
72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Length, 2.3—2.4 mm. Colour yellow-brown, the head a trifle darker. Hind legs,
especially the femora, dark brown.
Female. Head longer than wide across the eyes [10:9]; pubescent. Longitu-
dinal diameter of the eye three quarters the length of the cheek. Antenna (fig.
283): the scape large; the pedicel with several hundreds of small spines (fig. 285)
at the axial surface; the third segment rather robust, its appendage strongly curved.
Fourth segment small. Fifth segment almost glabrous, with ten oblong sensilla.
Sixth to ninth segments long, about twice the length of the fifth segment. Tenth
segment shorter; the apical segment but little longer than the fifth. Sixth to
eleventh segments with several irregular rows of sensilla. Labium with two sub-
apical hairs. Maxilla (fig. 291) with one subapical lateral hair, and a bacilliform
process, which is nearly half as long as the maxilla itself. The process bears one
apical hair, and a lateral hair at four-fifths of the length. The mandible (fig.
290) bears nine ventral ridges, its appendage seven.
Thorax. Pronotum rather long, with long hairs. Scutum about as long as its
maximal width; its lateral and ventral edges with about ten long hairs. Anterior
width of the scutellum five-sevenths of its length; the scutellum bears approxi-
mately forty long hairs. Metanotum with rows of fifteen hairs on each side of the
mid line. Propodeum long, with tufts of seven hairs above, and nearly fifty hairs
beneath and next to the spiracular peritremata. Fore wing (11 : 5), 3.0 mm long.
Submarginal, marginal, stigmal, and postmarginal veins approximately in ratio
16:5:4:11; the stigma distally produced, with two pairs of pustules; the sub-
marginal vein with three pustules. Hind wing (4:1), 1.85 mm long. Fore leg
(fig. 280): the coxa two-thirds, the tibia (including the dorsal armature) one half
the length of the femur. Tibia with a dorso-apical comb of four sharp and one
blunt teeth; ventrally with a blunt protuberance, which bears several long hairs.
Axially, the tibia and the tarsus bear several long, stout spine-like hairs. Tarsal
segments in ratio 18:5:5:4:8. Mid leg slender; the coxa semiglobular, the
trochanter short, but rather wide, the subclavate tibia nearly as long as trochanter
and femur combined, with one long ventral spur; tarsal segments in ratio 10 : 4:
4:3 : 6. Hind leg (fig. 282): the coxa as wide as the femur, but shorter [4 : 5};
the femur pubescent, with a distinct ventral groove for the reception of the tibia.
Tibia with two ventro-apical teeth, the antaxial one of which is bidentate. Tarsus
with spines and many hairs, especially along the plantar edge, the segments in
ratio ING) 89) 8 7/ AIS,
Gaster. Ovipositor short, approximately one tenth the length of the gaster.
Length, 3.2—3.4 mm. Colour of head and dorsal surfaces of thorax and gaster
black-brown, lighter ventrally. Nervures of wings brown, with dark striae radiating
from the stigma.
Remark.
C. pilipes is remarkable in having the dorso-apical edge of the male hind
tibia produced, and the parameres strongly curved. The dorsal armature of the
female fore tibia — and this holds for several other species in the relationship of
C. pilipes — reminds one of the tibial comb in C. armipes and relatives.
J. T. WIEBES : Indo-Australian Ceratosolen 73
SQ
N
SR \ 4 ZA
LE
Figs. 278—292, Ceratosolen pilipes (286, C. josephi), Borneo, 278, male mandible, 279,
male, 280, detail of female fore leg, 281, male antenna, 282, detail of female hind leg,
283, female antenna, 284, male, cercus of tenth urite, and paramere, 285, female, axial
spines of pedicel, 286, do., of C. josephi, 287—289, male, 287, hind leg, 288, labium and
maxillae, 289, metasternum, 290, female mandible, 291, female maxilla, 292, male fore leg.
Figs. 279, 287, 289, X 40; 280, 282, 283, 290—292, X 90: 278, 281, X 115; 285, 286,
288, X 200; 284, X 275
74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Ceratosolen josephi sp. n. (figs. 286, 293— 303)
Material.
Series 9, &, ex Ficus francisci Winkler (det. E. J. H. CORNER), Sungei Mamut
(N. Borneo), 3000 ft. alt, leg. E. J. H. Corner, VIII.1961; coll. no. 621; &
(holotype), slide 621a, 9 (allotype), 621c, 4, 9 (paratypes), 621b, d-g.
Dieisieir itp ition:
Male. Head (fig. 296) twice as long as wide anteriorly. The maximum width
is but little longer than the anterior width [5 : 4]. Pubescence short, especially
occurring along the antaxial edges of the antennal grooves. Eyes present. Antennal
grooves open, wide behind. Antenna (fig. 294) five-segmented; the scape, not
nearly twice as long as wide apically, twice as long as the pedicel (3 : 2); the
first flagellar segment annuliform, the apical segment (7:3) thrice as long as
the second flagellar segment (9 : 7). Mouthparts, fig. 301. Labium with a pair of
subapical hairs; maxilla expanded laterally, with two lateral hairs. Mandible, fig.
298.
Thorax, fig. 296. Pronotum twice as long as wide anteriorly, and distinctly
longer than wide posteriorly [6 : 5}. Maximum width of the mesonotum four-
fifths of its length. Metanotum incompletely separated from the propodeum,
which is but little longer than its maximum width. Metasternum with heavy pu-
bescence, mesosternum with less hairs. Fore leg (fig. 303): the femur more than
twice as long as the tibia [11 : 5}; the four dorsal teeth of the tibia not so close
together as in C. pilzpes. Tibia with three ventral apical teeth. Tarsus bimerous,
with notches indicating the presence of a third intermediate segment; the two
free segments subequal. Mid leg: the coxa smaller than the femur; the tibia smaller
than femur and trochanter combined. Tibia with long dorsal and ventral hairs, the
ventral edge produced into two apical teeth. Tarsus with long hairs, the segments
in ratio 9:4:4:4:8. Hind leg (fig. 299) with heavy pubescence on all seg-
ments. Coxa rather narrow, shorter than the femur, the tibia two-thirds the length
of the femur. Dorso-apical edge of the tibia produced; ventrally, the apical edge
bears a bidentate tooth, and a sharp tooth at the outermost corner. Tarsal segments
in ratio 9:7:6:4:7; pubescence nearly as long as the width of the segments.
Gaster partly sclerotized. Cerci of the tenth urite with three short claws, para-
meres almost straight (fig. 297).
Length, 2.0—2.2 mm. Colour light brown, with darker hind legs.
Female. Head as long as wide across the compound eyes; with rather long hairs.
Longitudinal diameter of the eye four-fifths of the length of the cheek. Antenna
(fig. 293): scape not very large relative to the large pedicel, which bears approx-
imately one hundred and fifty spines. Compared with those of C. pzlipes, these
spines are rather large (fig. 286). Fifth segment with ten oblong sensilla. Sixth to
eighth segments longer than the ninth {11:10}, which is twice as long as the
fifth. Tenth segment smaller than the ninth, the apical segment slightly longer
than the fifth. Sixth to eleventh segments bear several irregular rows of sensilla.
Labium and maxillae (fig. 302): the labium bears two subapical hairs; the max-
J. T. WieBEs: Indo-Australian Ceratosolen 75
illa bears one subapical hair and a bacilliform process, which is half as long as
the maxilla. The apical hair of the bacilliform process is visible in all specimens.
The presence of a lateral hair is indicated by a small pit, at four-fifths of the
length; I cannot find any specimens in which the hair is actually present. Mandible
(fig. 300) with eight ventral ridges, the appendage with seven.
Thorax. Relative proportions much as in C. pilipes, but some parts are distinctly
less pubescent: scutellum with four hairs along each lateral edge; metanotum with
rows of four to five hairs on each side of the mid line; propodeum with about
twenty-five hairs below the peritremata. Fore wing (21:10), 2.55 mm long.
Submarginal, marginal, stigmal, and postmarginal veins in ratio 20:7:8:15.
Stigma rather blunt distally, with three pustules; submarginal vein with three
pustules. Hind wing (4:1), 1.6 mm long. Fore leg (fig. 295): the coxa nearly
two-thirds the length of the femur, which is more than twice as long as the tibia
including the dorsal armature. Dorsal comb of the tibia consisting of four sharp
and one blunt teeth; ventrally, the tibia bears a protuberance with long hairs.
Axial surfaces of tibia and tarsus with several stout hairs. Tarsal segments in
ratio 22:7:8:7:14. Mid leg as in C. pzlipes. Femur of the hind leg rather
pubescent, twice as long as the tibia, which bears the usual axial, and bidentate
antaxial teeth. Tarsus rather wide, pubescent along the plantar edge; segments in
ZIO ORIO:
Gaster. Ovipositor distinctly projecting, one seventh the length of the gaster.
Length, 2.5—2.7 mm. Colour dark brown dorsally, lighter ventrally. Fore wings
with brown nervures and dark striae.
Remark.
This species is close to C. pzlipes, yet it is easily recognized by its smaller size,
and by several characters in the female antenna and thorax, and in the male mouth-
parts and legs.
I name the species after Dr. K. J. JosEPH, Head, Department of Zoology,
Karnatak University, Dharwar (India), in recognition of his contributions to the
knowledge of Indian fig insects.
Ceratosolen albulus sp. n. (figs. 304—316)
Material.
Series 9, &, ex Ficus uncinata Becc. var. strigosa Corner (det. E. J. H. Cor-
NER), Mt. Kinabalu East (N. Borneo), 3500 ft. alt, leg. E. J. H. CORNER,
VI.1961, RSNB no. 666; coll. no. 603; 4 (holotype), slide 603a, 9 (allotype),
603b, 9, 4 (paratypes), 603c, d.
Dieser pit von:
Male. Head (fig. 305) longer than its maximal width [5 : 4}, the anterior width
three-fifths of the length. Pubescence short and sparse. Eyes absent. Antennal
grooves open, wide behind. Antenna (fig. 306) five-segmented; the scape (3 : 2)
but little longer than the pedicel (4:1). First flagellar segment (1 : 2) one sixth
the length of the pedicel; the second segment nearly thrice as long as the first, and
one third the length of the apical segment (3 : 1). Labium and maxilla (fig. 316)
76 ‘TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Figs. 293—303, Ceratosolen josephi, Borneo, 293, female antenna, 294, male antenna, 295,
detail of female fore leg, 296—299, male, 296, male, 297, cercus of tenth urite, and para-
mere, 298, mandible, 299, hind leg, 300, female mandible, 301, male labium and maxillae,
302, female labium and maxilla, lateral aspect, 303, detail of male fore leg. Figs. 296, 299,
X 40; 293, 295, 300—303, X 90; 294, 298, X 115; 297, X 275
wide, with one lateral hair on the maxilla; labium bilobate. Mandible, fig. 315.
Thorax, fig. 305. Pronotum as long as wide posteriorly, the anterior width two-
thirds of the length. Mesonotum distinctly wider than long [17 : 14}, the lateral
edges with a stricture at mid length. Metanotum rather large, incompletely separated
from the propodeum; the latter longer than wide [5 : 4}, with straight lateral
margins. Fore leg (fig. 304): the coxa two-thirds the length of the femur, which
is twice as long as the tibia. Dorso-apical comb of the tibia consisting of three
large and two smaller teeth; there are several ventral teeth. Tarsus bimerous, seg-
ments approximately in ratio 2:3. Femur of the mid leg as long as coxa and
trochanter combined; the tibia slightly longer than femur and trochanter combined.
Ventral tibial edge produced into two apical teeth. Tarsal segments in ratio 17 :5 :
6:6:20. Femur of the hind leg (fig. 307) not much longer than coxa and tro-
chanter combined; the tibia distinctly shorter [4:5]. The coxa with a small
hyaline dorsal ridge, and many hairs on the axial surface; the femur and tibia, as
well as the tarsus, pubescent. Tibial armature consisting of two ventro-apical
J. T. WieBEs: Indo-Australian Ceratosolen zi
teeth (fig. 308), the antaxial one of which is bidentate. Tarsus dilated, the seg-
ments in ratio 45 : 22 : 23 : 20 : 40. Length of the pubescence one half to three
quarters the width of the segments.
Gaster. Cerci of the tenth urite with six claws; in many specimens, the para-
meres are folded longitudinally (fig. 309).
Length, 2.2—2.3 mm. Colour yellowish; the thoracic terga ivory-whitish.
Female. Head slightly shorter than wide across the eyes [17 : 18]. Pubescence
long. Longitudinal diameter of the eye longer than the cheek [5 : 3}. Antenna,
fig. 310. Scape two and a half times as long as the pedicel, which bears approxi-
mately fifty spines at the axial surface. Third segment small, not half as long as
the fifth; the latter two-thirds the length of the sixth; seventh to eleventh seg-
ments gradually diminishing in size, the eleventh about as long as the fifth,
but narrower. Sensilla rather long and narrow; fifth to tenth segments with about
twenty to twenty-five, the apical segment with fifteen sensilla. Labium and maxilla
(fig. 311): the labium with two hairs; the maxilla with a bacilliform process
(half as long as the maxilla) and a subapical, lateral hair. Mandible (fig. 312)
with seven ridges, the appendage with eleven ridges, the proximal two of which
are very small and not always distinctly visible.
Thorax. Pronotum with rather short hairs. Scutum anteriorly one and a half
times as wide as long, the posterior width half the anterior width; with five long
hairs along each lateral margin. Posterior width of the scutellum nearly equal to
its length, anterior width four-fifths of the length. Lateral margins with five
hairs. Metanotum rather long, with fourteen hairs on each side of the mid line.
Propodeum with two hairs above, and nine hairs beneath the peritremata. Fore
wing (2:1), 2.5 mm long. Submarginal, marginal, stigmal, and postmarginal
veins in ratio 39 : 21 : 20 : 25; submarginal vein with three pustules, stigma with
four. Hind wing (5:1), 1.6 mm long. Fore leg (fig. 314): the coxa more than
half as long as the femur {5 : 8}, which bears long hairs. Tibia, not nearly half
as long as the femur [3 : 8], with a dorso-apical comb of six teeth, the first of
which is blunt. Ventrally, the tibia bears a blunt tooth and a long spur.
Antaxial surface with long hairs, especially in the ventral part; axial surface with
ten stout spine-like hairs. Tarsal segments in ratio 13 : 5 : 5 : 6 : 7; axial surfaces
of the segments with stout spines, except for the fifth, which has hairs instead.
Mid leg slender, the tibia nearly as long as trochanter and femur combined; tarsal
segments in ratio 23 : 11 :13 :12 : 15. Hind leg (fig. 313); the femur slender,
one and a half times as long as the tibia. Tibia with long hairs and two ventral
teeth, the antaxial one of which is bidentate. Tarsus pubescent, segments in ratio
PONSO SIAE
Gaster. Ovipositor short, but distinctly protruding behind the gaster.
Length, 2.3—3.0 mm. Colour brown, pronotum and head darker, ventral sur-
face and extremities lighter. Fore wing with two dark striae radiating from the
stigma
Remark.
C. albulus is close to C. humatus, but it differs in its dimensions, the dentation
of the male fore tibia, the length of the pedicel of the male antenna, etc.
78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Ceratosolen humatus sp. n. (figs. 317—329)
Material.
Series 9, 4, ex Ficus beccarii King var. latifolia Corner (det. E. J. H. Cor-
NER), Kundasan (N. Borneo), 4000 ft. alt, leg. E. J. H. CORNER, 24.VIII.1961;
coll. no. 608; & (holotype), slide 608a, «9 (allotype), 608c, 4 (paratype),
608b.
Series 9, &, ex Ficus beccarii King var. latifolia Corner (det. E. J. H. Cor-
NER), Mt. Kinabalu East (N. Borneo), 6500 ft. alt., leg. E. J. H. Corner,
7.VIII.1961, RSNB no. 709; coll. no. 611; 9, & (paratypes), slides 611a, b.
Series 9, 4, ex Ficus beccarii King var. latifolia Corner (det. E. J. H. Cor-
NER), Mt. Kinabalu East (N. Borneo), 3500 ft. alt, leg. E. J. H. CORNER,
VI.1961, RSNB no. 665; coll. no. 614; ®, & (paratypes), slides 614a, b.
Series 9, 4, ex Ficus subterranea Corner (det. E. J. H. CORNER), Tenompok
(N. Borneo), 4500 ft. alt., leg. E. J. H. CORNER, VIII.1961, RSNB no. 1987; coll.
mo, COWS Slide COTE, 1, Oo 07 Gh
Description.
Male. Head (fig. 326) one and a half times as long as wide anteriorly, and
distinctly longer than its maximum width [9 : 7}. Pubescence short and sparse.
Eyes absent. Antennal grooves open, wide behind. Antenna (fig. 323) five-
segmented: the scape (12:7) twice as long as the pedicel (2:1); the first
flagellar segment (3 : 7) one third the length of the pedicel, the second segment
(2 : 3) twice, the apical segment (3 : 1) seven times as long as the first. Labium
and maxillae, fig. 319. Labium bilobate at apex, each lobe with an apical hair, and
sometimes with an extra hair at the axial surface; maxilla with two lateral hairs
as in the figure. Mandible, fig. 324.
Thorax, fig. 326. Pronotum short, as long as wide at mid length, slightly nar-
rower in front, and much wider behind. Mesonotum large, nearly as long as wide
posteriorly, narrower anteriorly. Metanotum incompletely separated from the
propodeum, the free part of which is about as long as wide. Coxa of fore leg
(fig. 325) large, more than half as long as the femur; the femur twice as long as
the tibia. Apical armature of the tibia consisting of a series of dorsal teeth, and three
ventral teeth. Tibia axially as well as antaxially with stout hairs. Tarsus bimerous,
although a dorsal notch in the distal segment indicates an original trimerous con-
dition; the free segments subequal in length. Mid leg slender, the coxa slightly
longer than the femur, but shorter than the clavate tibia. Tibia with long hairs;
its ventro-apical edge produced into two teeth. Tarsal segments in ratio 14:5 : 5 :
4:14, Hind leg (fig. 321): the coxa but little shorter than the femur, glabrous
antaxially, strongly pubescent axially, with a dorsal hyaline ridge. Femur antaxially
with few short hairs, pubescent axially. Tibia three quarters the length of the femur,
pubescent; the ventral armature consisting of a bidentate antaxial, and an axial
tooth, which is slightly curved. Tarsus dilated, the pubescence approximately one
half to five-sevenths the width of the segments. Tarsal segments in ratio 23 : 11 :
11:11:18, or the first segment slightly shorter and the tarsus wider, or the first
segment slightly longer and the tarsus longer, in some specimens.
J. T. WIEBES : Indo-Australian Ceratosolen 79
Figs. 304— 316, Ceratosolen albulus, Borneo, 304—309, male, 304, detail of fore leg, 305,
male, 306, antenna, 307, hind leg, 308, apex of hind tibia, 309, cercus of tenth urite, and
paramere, 310—314, female, 310, antenna, 311, labium and maxilla, 312, mandible, 313,
detail of hind leg, 314, detail of fore leg, 315, male mandible, 316, male labium and
maxilla. Figs. 305, 307, X 40; 310, X 90; 304, 306, 308, 311— 314, X 115; 315, X 160;
ON Sl 27/5
80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Gaster. Cercus and paramere, fig. 322; the cercus apically with four claws.
Apodemae aedeagales rather darkly sclerotized.
Length, 1.5—1.9 mm. Colour uniformly whitish yellow, darker in older speci-
mens.
Female. Head short, not quite as long as wide across the eyes [23 : 24].
Pubescence long. Longitudinal diameter of the eye twice as long as the cheek.
Antenna (fig. 318): the scape thrice as long as the pedicel, which bears ap-
proximately forty spines. Appendage of the third segment slender, rather long;
the fourth segment small. Fifth segment twice as long as the fourth, with twelve
long sensilla. Sixth and seventh segments one and a half times as long as the
fifth, but narrower; eighth to eleventh segments gradually diminishing in size.
Sixth to tenth segments with irregular, alternating rows of long sensilla: ap-
proximately twenty sensilla per segment; apical segment with twelve long and two
circular sensilla. Labium with two subapical hairs. Maxilla (fig. 317) with one
subapical lateral hair, and a bacilliform process, which is more than half as long
as the maxilla. Mandible (fig. 320) with six ventral ridges, its appendage with
nine.
Thorax. Pubescence of pronotum long. Scutum shorter than its maximal width,
with six hairs along each lateral margin. Scutellum one and a half times as long
as wide, its sides (except posteriorly) subparallel, with five or six hairs along each
lateral edge. Metanotum with eleven hairs on each side of the mid line; propodeum
with three hairs above, and about ten beneath the large peritremata. Fore wing
(2:1), 2.1 mm long; submarginal, marginal, stigmal, and postmarginal veins in
ratio 8:4:3:5; submarginal vein with two pustules, stigma with four. Hind
wing (5:1), 1.4 mm long. Fore leg (figs. 327—328): the coxa two-thirds the
length of the femur; the tibia not quite half as long as the femur. Tibial armature
consisting of a dorsal comb of six teeth (the first blunt), and one ventral tooth.
Axially, the tibia is provided with several stout hairs. Tarsal segments in ratio
11:5:5:5:7. Mid leg slender, the coxa semiglobular; the tibia not quite as
long as femur and trochanter combined. Ventral edge of the tibia produced apical-
ly. Tarsal segments in ratio 25 :10:9:8:12. Coxa of the hind leg (fig. 329)
shorter than the femur, but wider; the femur narrow; the tibia slender, two-thirds
the length of the femur. Tibial apex axially with a curved tooth, antaxially with
a bidentate tooth. Tarsus pubescent, the segments in ratio 25:11:11:9:10.
Gaster. Ovipositor scarcely projecting beyond the apex of the gaster.
Length, 2.2 mm. Colour blackish brown dorsally, lighter ventrally. Extremities
light yellow-brown. Old specimens are darker; young specimens sometimes have the
last two or three antennal segments of an ivory-white colour. Fore wings with two
dark striae radiating from the stigma.
Remark.
C. humatus is evidently related to C. pilipes and C. josephi, but the male meta-
sternum is glabrous, and the hind tibia is not produced dorsally; the female tibial
comb consists of six teeth.
I cannot find distinct constant differences between the wasps from Ficus bec-
nr—_——————F6——__n
J. T. WieBEs: Indo-Australian Ceratosolen 81
carit and F. subterranea. Although the specimens from coll. no. 608 have the male
hind legs slightly more slender than those of coll. no. 607, this differential char-
acter is bridged by the males from coll. no. 611. Consequently, I refer the spec-
imens from F. subterranea to C. humatus.
Ceratosolen bisulcatus (Mayr) and C. jucundus Grandi (figs. 152, 153;
map, fig. 261)
C. bisulcatus (Mayr) and C. jucundus Grandi are so similar that it is question-
able whether the two should be maintained as separate species. Some differences
between C. bisulcatus from Java and Bali, and C. jucundus from Luzon, are:
9. Maxilla with a bacilliform process in C. j#cundus, without such a process in
C. bisulcatus (fig. 153).
4 . The head is slender in C. bisulcatus (one and a half times as long as wide),
and bears small eyes. In C. j#cundus the head is relatively shorter, and the form
is anophthalmous.
Specimens from New Guinea are slightly longer than those from Java and Bali,
and the females are darker. Specimens from Borneo are of the general facies of
C. jucundus from Luzon, but the female maxilla does not bear a bacilliform pro-
cess, or, at most, it has a small protuberance instead (in this it is similar to the
form recorded from Formosa, of which I did not see examples). Distribution,
see map, fig. 261. The male mouthparts (fig. 152) and genitalia (cercus of the
tenth urite with two claws) are similar in the two species, and both show the
oligomery in the male mid tarsi (but these are pentamerous in the specimens
from Formosa, and heteromerous in those from Borneo). C. bisulcatus males
have heteromerous hind tarsi, which I do not find in C. jucundus.
Ceratosolen bisulcatus (Mayr)
Blastophaga (Ceratosolen) bisulcata Mayr, 1885, pp. 154, 161, 163, 170—171 [key ®, 4,
descr. 9, 8, ex Ficus (Cystogyne) lepicarpa BI. (det. Solms)!), Bogor (Java), Bot. Gdn.;
type: à |.
Ceratosolen bisulcatus: Karny, 1923, figs. 5—6 [@, &, Krakatau, leg. K. W. Dammer-
man}; Grandi, 1928c, pp. 180—184, figs. XXIX—XXX [redescr. 9, 4, type specimens
of B. bisulcata Mayr}; Dammerman, 1948, pp. 381—382, fig. 21 [®, &, Krakatau, and
Verlaten Island, IX.1920, 1933}.
Material.
Series 9, 4, ex Ficus septica Burm. f., Bogor (Java), Baranangsiang, 24.X.
1954; coll. no. 75.
Series 9, &, ex Ficus septica Burm. f. (det. E. J. H. CORNER), Lae (Terr.
New Guinea), Erap, leg. E. J. H. CORNER, 7.IX.1960; coll. no. 544; &, slide
544a, 9, 544b.
Series 9, 4, ex Ficus septica Burm. f., Sudadji (Bali, distr. Sawahan, 16 km
SE of Singaradja), leg. NJOMAN ToJA; coll. no. 58; &, slide 58a.
Series 9, 4, ex Ficus spec., Java, leg. A. HOOGERWERF, X1.1954, no. 9; coll.
no. 283.
1) SoLMs evidently confused F. septica with F. lepicarpa; see p. 98.
82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Figs. 317—329, Ceratosolen humatus, Borneo, 317, female maxilla, 318, female antenna,
319, male labium and maxillae, 320, female mandible, 321—326, male, 321, detail of hind
leg, 322, cercus of tenth urite, and paramere, 323, antenna, 324, mandible, 325, detail of
fore leg, 326, male, 327— 329, female, 327, detail of fore leg, axial aspect, 328, fore tibia,
329, detail of hind leg. Figs. 326, X 40; 318, 321, 329, X 90; 317, 320, 323—325, 327,
DAR XX LE IAD X 275
J. T. WieBEs: Indo-Australian Ceratosolen 83
Series 2, 4, ex Ficus septica Burm. f., Udjung Kulon (Java), leg. A. HOOGER-
WERF, XI.1954, no. 11; coll. no. 273; 9, &, slide 273a, immature specimens,
273b.
Series 9, &, ex Ficus septica Burm. f., Udjung Kulon (Java), leg. A. HOOGER-
WERF, XI.1954, no. 16; coll. no. 287; ®, slide 287a, 4, 287b.
Ceratosolen jucundus Grandi
Ceratosolen jucundus Grandi, 1927a, pp. 320—323, PI. 5 figs. 67—76, PI. 6 figs. 77—83
[descr. 2, &, ex Ficus hauili Blanco!), Mt. Maquiling (Luzon, Philippine Is.), leg. C. F.
Baker}; Grandi, 1927b, pp. 178-179 [descr. 9, 8, C. ? jucundus, Taihoku (Formosa),
leg. F. Silvestri, 21.X1.1954}.
Material.
Three 9, 3 &, ex Ficus hauili Blanco!), Mt. Maquiling (Luzon, Philippine
Is.), ex coll. GRANDI; coll. ML, no. 507.
Series ?, &, Mt. Maquiling (Luzon, Philippine Is.); coll. USNM.
Series 9, 4, ex Ficus hauili Blancot), Los Baños (Luzon, Philippine Is.),
Coll. Agric., leg. F. X. WILLIAMS, 6.II, 21.V, and 19.VI.1921; coll. HSPA.
Series 9, 4, ex Ficus hauli Blanco!), Lake Lanao (Mindanao, Philippine Is.),
leg. F. X. WILLIAMS, 2.X1.1921; coll. HSPA; coll. ML, no. 587; ®, slide 587a,
4 Sib:
Series 9, &, ex Ficus septica Burm. f., (det. E. J. H. CORNER), Ranau (N.
Borneo), 1800 ft. alt., leg. E. J. H. CORNER, 3.VI.1961; coll. no. 616.
Series 2, 6, ex Ficus septica Burm. f. (det. E. J. H. CORNER), Kundasan (N.
Borneo), 4000 ft. alt., leg. E. J. H. CORNER, IX.1961; coll. no. 634.
Species incertae sedis
Ceratosolen spec.
Material.
Immature 9, &, ex Ficus microdictya Diels (det. E. J. H. CORNER), Papua,
leg. C. E. Carr, 14.X11.1935, no. 13785; coll. no. 449.
Remark.
From Ficus microdictya, I saw some immature wasps only, which I prefer not
to describe. They seem to be close to C. armipes and allied species.
Ceratosolen spec.
Ceratosolen ? crassitarsus: Grandi, 1923a, p. 299 [descr. note on 92, 6, ex Ficus
roxburgh®), Singapore, leg. H. N. Ridley}; 1927b, p. 174 {note on host record].
Material.
One 9,5 4, ex Ficus Roxburghii2), Singapore, H. N. RIDLEY, 89—38; coll.
BM: 5 slides.
1) = Ficus septica Burm. f.
2) Synonym of Ficus auriculata Lour., but the record is probably incorrect.
84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Remark.
In my opinion these specimens should be referred to a form of Ceratosolen
vechti, rather than to C. crassitarsus; but the description has to wait until more
and better material becomes available.
Ceratosolen s pec.
Material.
Fragments of 3 9, ex Ficus stolonifera King (det. E. J. H. CORNER), Sarawak
(Borneo), leg. M. JACOBS, no. 5257; coll. no. 503.
Remark.
These specimens, taken from dried figs in the Leiden Herbarium, are too frag-
mentary for specific identification, but they can be recognized as belonging to a
species of Ceratosolen. The dorsal comb of the female fore tibia consists of four
sharp teeth, and it does not show the blunt tooth found in other Ceratosolen from
geocarpic figs.
Provisionally excluded
Ceratosolen megarhopalus Grandi, 1923b, pp. 103—104 [descr. 9, at light,
Fort de Kock (Sumatra), leg. E. JACOBSON, X1.1920, 1.1923}.
Ceratosolen elisabethae Grandi, 1923b, pp. 104—105 [descr. 9 , at light, Fort
de Kock (Sumatra), leg. E. JACOBSON, XI.1920, I. 1923].
These two species, which are known in the female sex only, are characterized
by the peculiar antennae, and the large eyes.
In the collection made by Dr. F. X. WırLıams in the Philippine Islands, I
find a sample of females and males ex Ficus ? longipedunculata, Mt. Maquiling
(Luzon), 23/24.VI.1921; coll. HSPA, which evidently belong to a species close
to C. megarhopalus Grandi. The males have a very distinct Blastophaga facies, and
this is the reason why I exclude the species of this group from the present revision
of Ceratosolen.
Other samples ex “Frcus longipedunculata’ (without the query; F. longipe-
dunculata (Merril) Elmer = F. chrysolepis Miq., subgenus Urostigma) consist
of a different species of Agaonidae.
CLASSIFICATION AND KEY TO THE SPECIES
In this chapter an attempt is made to classify the species in groups. I am well
aware that these groups must be considered provisional divisions, which show some
overlapping.
It is difficult to arrive at a satisfactory conclusion as to the status of some taxa.
In one instance (C. solmsi and C. marchali, cf. p. 64, Table I) it is evident that
the two forms should be regarded as subspecies. In other instances this evidence is
lacking or non-conclusive, due to the scantiness of the material. For the time
being, pending the examination of more material from more localities, I treat the
J. T. WieBES: Indo-Australian Ceratosolen 85
following forms: C. constrictus and C. hewitti; C. notus, C. orientalis, and C.
calopilinae; C. bisulcatus and C. jucundus, as full species.
I. C. pygmaeus group.
In this group I place C. pygmaeus, C. marshallt, and C. nanus, which all have
the antennal grooves in the male head half closed, the labium and the maxillae
atrophied, the lateral edges of the propodeum more or less rounded, the female
maxillae without a bacilliform process, and the fore tibia with no more than three
teeth in the dorsal comb and without the ventral spur.
C. nanus is aberrant in some aspects (male antenna consisting of five segments,
genitalia without cerci, female antenna of a peculiar shape, its segments with few
sensilla), but the trimerous mid and hind tarsi in the male, and the tetramerous
fore and mid tarsi in the female, next to the general facies, connect the species
with C. pygmaeus.
C. gravelyi, although evidently related to the species of this group (facies,
male tibia without spines on the disk), seems to show more connections with the
C. appendiculatus group.
I have long hesitated about the classification of C. constrictus and C. hewitti,
which show some relationships with the C. pygmaeus group in the general facies,
the oligomery of the mid and hind tarsi, and in the absence of the ventral spur
from the female fore tibia. On the other hand, they have four teeth in the combs
of the male and female fore tibiae (except for the sample from N. Borneo), as in
some other groups. Tentatively I place C. constrictus and C. hewitti in this C.
pygmaeus group.
Species of the C. pygmaeus group:
C. pygmaeus Grandi, C. constrictus (Mayr),
C. marshalli Grandi, C. hewitti Waterston.
C. nanus Wiebes,
II. C. appendiculatus group.
The species of this group have the antennal grooves in the male head half
closed; as a rule the mid and hind legs bear spines on the disks, and the shape
of the propodeum is rather uniform. The female fore tibia bears a simple ventral
spur, and the dorsal dentation consists of four sharp teeth.
C. gravelyi connects the C. pygmaeus group with C. emarginatus, which at its
turn resembles C. fusciceps and C. appendiculatus. Apparently C. appendiculatus
and C. grandii are closely related: both species have the male genitalia without
cerci and the club of the female antenna is composed of the united apical three
segments.
Species of the C. appendiculatus group:
C. gravelyi Grandi, C. appendiculatus (Mayr),
C. emarginatus Mayr, C. grandii Wiebes.
C. fusciceps (Mayr),
Judging from the descriptions, the African species of Ceratosolen belong in
this group.
86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
III. C. armipes group.
C. armipes, C. bianchii, C. boschmai, and C. sordidus have in common the
half closed antennal groove, the spines on the disk of the mid tibia, and the
aberrant shape of the propodeum of the male, the dentation of the comb of the
female fore tibia, and to some extent the general facies.
The group in which these species are united shows a rather great variation, e.g.,
in the shape of the male propodeum. In a way, the species resemble C. pilipes
and its relatives, notably in the dentation of the female fore tibia.
Species of the C. armipes group:
C. boschmai Wiebes, C. armipes Wiebes,
C. bianchii Wiebes, C. sordidus Wiebes.
IV. C. bakeri group.
Some characters connect C. bakeri with the C. armipes group (male antenna
resembling that of C. boschmai, male propodeum, dentation of female hind tibia),
but the species has open antennal grooves and a three-segmented fore tarsus in
the male.
Species of the C. bakeri group:
C. bakeri Grandi.
V. C. abnormis group.
C. abnormis, though evidently related to the species of the C. armipes group in
some aspects (male antennal grooves half closed, segments of the fore tarsus in-
completely separated, dentation of the female hind tibia), is aberrant in the
male genitalia and in the armature of the female fore tibia.
It is classified here as a separate group:
C. abnormis Wiebes.
VI. C. crassitarsus group.
Characterized by the dilated hind feet of the male, the normal mouthparts —
without large lateral expansions on the maxillae — and the dentation (four
sharp teeth) of the comb of the female fore tibia. In some species (C. 72045,
C. moderatus, etc.) the male maxillae bear small lateral expansions, thus con-
necting the group with the C. pilipes group. Some other species are aberrant in
the dentation of the male hind tibia, e.g., C. nugatorius and C. dentifer have a
large motile spur on the disk of the hind tibia; C. corneri is aberrant in having
very long hind feet in the male sex. Otherwise, the species of this group are very
similar, and it is difficult to distinguish between the females of the various
subgroups.
Species of the C. crassttarsus group:
. crassitarsus (Mayr),
. gracilis Wiebes,
. hooglandi Wiebes,
@ . praestans Wiebes,
G
G
C. vechti Wiebes,
G
C
C
iodotrichae Wiebes,
notus (Baker),
. orientalis Wiebes,
calopilinae Wiebes,
corneri Wiebes.
. nugatorius Grandi,
. dentifer Wiebes,
. moderatus Wiebes,
ADASAG
J. T. Wiegers: Indo-Australian Ceratosolen 87
VII. C. solmsi group.
Two species with normal hind feet in the male (others with the same character,
viz., C. bisulcatus and C. jucundus are aberrant in other features too and are joined
to represent a distinct group IX) are difficult to be classified.
C. solmsi is evidently related to species with dilated hind feet, notably the
C. crassitarsus group. In some characters, however, it is close to C. brongersmai
(e.g., in the dentation of the male hind tibia) and in the dentation of the female
fore tibia it differs from both the C. crassitarsus group and C. brongersmai.
C. brongersmai seems to be close to C. albulus and C. humatus, although it has
the hind feet of the male not dilated and the comb of the female fore tibia is of
normal constitution.
For convenience’ sake the two species are here taken together in one group,
which connects the C. crassitarsus group with the C. pilipes group.
The occurrence of C. brongersmai and C. solmsi among the species with dilated
hind feet in the male makes it plausible that the character “dilated hind feet”
may have been more than once acquired, or secondarily lost, in the evolution of
the group.
Species of the C. solmsi group:
C. solmsi (Mayr), and subspec. marchali Mayr,
C. brongersmai Wiebes.
VIII. C. pilipes group.
Some species have the male maxilla with a very large, lateral expansion, and the
female fore tibia with a blunt first tooth in the dorsal comb. C. pzlipes and C.
josephi have the metasternum of the male hirsute, and the apical edge of the
hind tibia produced dorsally; the fore tarsus shows three incompletely separated
segments. The tibial comb of the female fore leg consists of five teeth.
C. albulus and C. humatus have a glabrous metasternum, and the male hind tibia
is not produced. The combs of the fore tibiae consist of six teeth in the females,
and of five and nine teeth in the males of C. albulus and C. humatus, respectively.
Species of the C. pilipes group:
C. pilipes Wiebes, C. albulus Wiebes,
C. josephi Wiebes, C. humatus Wiebes.
IX. C. bisulcatus group.
C. bisulcatus and C. jucundus cannot easily find a place in any of the other
groups. Apparently they are close to the species of the C. crassitarsus group; but
they differ from all other species of Ceratosolen in the bilobate epistomal margin
of the male.
Species of C. bisulcatus group:
C. bisulcatus (Mayr),
C. jucundus Grandi.
KEW EO Ine Species
But for C. gravelyi, which is intermediate between the C. appendiculatus group
and the C. pygmaeus group, and keys out with the latter although classified in
88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
the former, all species groups can be identified by using the key, provided male |
specimens are available.
The following remarks may be helpful for identifying female specimens. Each
alternative excludes the preceding one(s).
1. If the fore tibia is without a ventral spur, try no. 4 (C. pygmaeus group),
and compare C. abnormis and C. bakeri.
2. If the comb of the fore tibia consists of more than four teeth, the first of |
which is blunt, try nos. 13 (C. armipes group) and 34 (C. pilipes group).
3. If the apical segments of the antenna are united (te, the antenna consists of |
less than eleven segments), try no. 10 (some species of the C. appendiculatus
group). |
4. If the apical three segments of the antenna are shaped so as to form together
a club, compare C. gravely: and C. emarginatus.
5. If the apical two segments of the antenna are shaped so as to form together a
(sometimes very loose) club, try no. 18 (C. bisulcatus and C. solmsi groups), and
compare the species of the C. crassitarsus group. The latter are not keyed out; it
will be possible to arrive at a satisfactory identification only after careful com-
parison with the descriptions.
nde Antennalsoroovesmhalfclosedm(Exam ple: fig: 20) DE?
— &. Antennal grooves open (example, fig. 180) . . . CIO
2. 4. Propodeum with comparatively small spiracular atrata 2. Fore
tibia with tworto foun sharp teethiin the dorsal comb 5
— 4. Peritremata of the propodeum large. ©. Fore tibia with five to ten teeth
in the dorsal comb. . BUND
3. &. Mid and hind tibiae han ines on he distel G premaens Bean
and C. gravelyi) . ue
— &. Mid tibia, and mostly the Rind de ale. Sir many ins on dn
disk (C. appendiculatus group except Go GASP) mea 9
4. &. Genitalia without cerci. 9. Antenna aberrant (fig. 16), sath fe sen-
SIA EN sn Blin SE GENT
— 4. Genitalia ul Gerci. 9). Aden oro, thank segments with one
or two rows of sensilla. . . . SS
5. &. Antenna consisting of four eames Labor ae zesde atrophied.
0%, Forertibia withrthree teeth:in the dorsal Comb mn
— 4. Antenna consisting of five segments, one of which may be annuliform.
Mouthparts present. @. Fore tibia with two or four teeth in the dorsal
combien: 7
6. &. Propodeum with len rounded alga Ose of the en urite with
one claw. 9. Mandible with ten ventral ridges, its appendage with nine
Eid ges) war Te EG. pygmaeus
— 4. Date more alles behind Ge 10). Cad with three claws. 9.
Mandible, and its appendage, with four or five ventral ridges . C. marshalli
7. 4. Thorax constricted (figs. 1, 2). 9. Funicular segments with two to three
rows of sensilla. Maxilla without a bacilliform process. Fore tibia without a
ventra lis pied Mund rt de... te NER ARR
I:
13:
J. T. WIEBES : Indo-Australian Ceratosolen 89
&. Thorax not as above. 9. Funicular segments with one row of sensilla.
Maxilla with a bacilliform process. Fore tibia with a ventral i ! i
Ean
Di Bed ad one Tal: shone robust Ge 2). 9. Sixth cn segment
as long as the seventh, with three rows of short sensilla; the tenth segment
approximatelyas#longasithereléventh rn. ON ON Gi constrictus
&. Head and thorax more slender (fig. 1). 9. Sixth antennal segment two-
thirds the length of the seventh, with two irregular rows of longer sensilla;
the tenth segment BA one and a half times as long as the
eleventh. . . . ; NT at Go MAIN
&. Mid and hind ee th ines or obees on the disks. 9.
Apical two or three antennal en united. Maxilla without a bacilliform
Process er: . 10
&. Hind tibia ae es on the det on Antennal Lunari frees but
the apical three segments shaped so as to form together a distinct club. Maxilla
with a bacilliform process. . . ee Cr ROD
&. Antenna consisting of five armen, Can and maxillae atrophied.
g. Funicular segments with two rows of sensilla. . . C. appendiculatus
8. Antenna consisting of four segments. Mouthparts present. 9. Funicular
segments with one row of sensilla. . . . . Poi ea
&. Tarsal segments of the fore leg incompletely separate Ge without
cerci. 9. Apical three antennal segments united. . . . . C. grandi
&. Tarsal segments of the fore leg free. Genitalia with cerci. 9. Apical two
antennalisegments united mann tl: NEGEN
4. Tibiae without spines on the disks. Genitals with parameres only (fig.
81). 9. Antenna with very few sensilla. Comb of the fore tibia consisting of
cightatomtenmshanpy teeth. 2. seb ls be Gy CGO
&. Mid and hind tibiae with spines or stout ne on the disks. Genitalia not
as above. 9. Funicular segments with one to three rows of sensilla (unknown
in one species). Comb of the fore tibia consisting of five or six teeth, the
first of which is blunt (C. armipes group). . . Mapa een al!
é. Antenna consisting of four segments. Fore tibia ah ce dorso-apical
teeth. 9. Apical two antennal segments united. . . . . . C. armipes
&. Antenna consisting of five segments, one of which may be annuliform,
or incompletely separated. Fore tibia with four dorso-apical teeth. 9 . Antennal
segments free, although the apical three shaped so as to form a club. . 14
4. Tarsal segments of the fore leg free. 9. Comb of the fore tibia con-
SiStING) OfMsixe teeth I gens aen (Ge WOM
4. Segments of the fore tarsus incompletely separated, g. Fore tibia with
fiverteethminmtherdorsalkcon bas MCE a EEA REEN U)
4. Genitalia with cerci. 9. Funicular segments with one row of sensilla
Antaxial apical tooth of the hind tibia bidentate. . . . . C. boschmat
4. Genitalia without cerci. 9. Funicular segments with two rows of sen-
silla. Antaxial apical tooth of the hind tibia tridentate. . . C. bianchi
Boele mel staet moek not eel dre lint enor tiny Ac ende oee A
Awaltindsteetwatlatedrandehirsutemes. RENTE n EERE 21
90
18.
12?
20.
21.
23:
24.
25
26.
27e
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
8. Antenna consisting of six segments, one of which is annuliform. Fore
leg with three tarsal segments. 9. Antaxial apical tooth of the hind tibia
EEn taten rn: eN PARENT
8. Antenna concern of fe sammen, one ab een may be annuliform.
Tarsus of the fore leg consisting of two free segments. 9. Antaxial apical
tooth of the hind tibia bidentate. . . LOS
&. Epistomal margin bilobate. 9. Eene sements ich one row a sen-
silla (C. bisulcatus group) . . SARTO)
&. Epistomal margin trilobate. 9. Banier nen two or three
rows of sensilla (C. solmsi group) . . HE R20
&. Head one and a half times as Lome = as ide - yes piane a. Maxilla
without a bacilliform process. . . a (Co LANG
d. Head relatively shorter, the lea not more (han four thirds of the
maximum width. Eyes absent. @. Maxilla mostly with a long bacilliform
process, or with a small protuberance instead. . . . . . €. jucundus
&. Maxillae with lateral a ©. Fore tibia with four teeth in the
dorsalecombrr rs De EEE AC brongersmai
4. Maxillae without a Sung a. “Fate tibia with five teeth in the
dorsal COM Eede vol Cs GOD
3. Propodeum wider than onz or taie the none ee angles rounded.
g. Comb of the fore tibia consisting of four sharp teeth (C. crassitarsus
group) . ANNEE 22
du Prg odaum urge: ion i ‘the eastern anes slightly produced
laterally. 9. Comb of the fore tibia consisting of five or six teeth, the first
Ofawhichpisiblunta(Geoz pes oro) EIA EEE RD
Ss HindWleszverAlongFasan®t19 SAN RE nin CGO nere
dinde noel mn DS RE 23
4. Hind tibia with a large dark pee ice 144, 157) « ne: os ee
&. Hind tibia with the normal apical armature only. . 5:25)
3. The spur on the hind tibia is situated apically. 9. RR seamen
with one row of very ee sensilla. Maxilla with a bacilliform process .
Ce Dia
a Te Spar is ne, more ce pendel © on the die 9. Funicular segments
with two rows of normal sensilla. Maxilla without a bacilliform process .
Halen ee Roe C. nugatorius
&. Hind tibia approximately twice as long as the metatarsus. . . . . 26
&. Hind tibia distinctly less than twice as long as the metatarsus . . . 27
4. Maxillae with nearly straight lateral edges, without any lateral expansions.
ua of male and female, os 1 and 1.5 mm, respectively .
la C. crassitarsus
8: Maxillae eze äh had npe Length o of male and female, ap-
proximately 1.5 and 2 mm, respectively . . ae 8 Go MOUS
8. Hind tibia as long as or up to one and a half times as long as the meta-
tarsus ENS 5 2S
4. Hind na porno « one emd Bees etes ie lea on the meta-
tarsus ae „ev Val ne apa ea a e AC
Bile
DE
99:
34.
DD:
36.
J. T. WieBES: Indo-Australian Ceratosolen 91
&. Length of the apical antennal segment one and a half to one and three
quarters the length of the subapical segment. . . . 1,29)
4. Apical segment of the antenna approximately two a a half times as
long as the subapical one, or longer . . . 50
&. Maxilla with a lateral expansion. Cerci gi {he (no urite sf five
Clas oo de (Oer VOOG)
&. Maxilla nent Tate En Gani of the tenth urite with two
Claws , nono mo. Go, (UV
a Gerci of e ‘i urite o seven alae. on Bacilliform process one
fifth the length of the maxilla. . . . ers ve Gn ANA
&. Cerci of the tenth urite with three to fe Ee 9. Bacilliform process
nearly half as long as the maxilla. . . . MAPS MT
8. Apical segment of the antenna two and a a times as one as the sab
apicaliseomenti IN : ee CN 00 landi
4. Apical segment of the antenna six times as one as the subapical segment
C. sodotrichae
a al geren di tho! antenna ‘Tess an twice as long as the subapical
segment. 9 . Bacilliform process one third the of the maxilla . 3
a 3 C. vechti
Si. Apical onen! of the antenna roten two and a half times as
long as the subapical segment. ©. Bacilliform process two-fifths the length
of the maxilla, or longer . . . . 33
&. Maxilla with two lateral hairs. Hind EE thee ea ese gi Maxilla not
quite twice as long as its bacilliform process. . . . . . C. orientalis
4. Maxilla with one lateral hair. Hind leg shorter, more robust. 9. Maxilla
ia two and a half times as long as the bacilliform process .
; 5 5 C. calopilinae
on BOC tarsus tore se ee) sent de distal two of which are
incompletely separated. Dorsal edge of the hind tibia produced apically.
Metasternum hirsute. 9. Comb of the fore tibia consisting of five teeth . 35
4. Fore tarsus consisting of two free segments. Dorsal edge of the hind tibia
not produced apically. Metasternum not hirsute. 9. Comb of the fore tibia
Consisting sofmstxeitecthiys A En ae ee
&. Cerci of the tenth urite with ne or six ek parameres curved. 9.
Pedicel of the antenna with several hundreds of small spines. . C. pilipes
6. Cerci of the tenth urite with three claws, parameres straight. 9 . Pedicel
of the antenna with approximately one hundred and fifty spines. . C. josephi
&. Fore tibia with a dorsal comb of five teeth. Pedicel of the antenna nearly
as long as the scape. Cerci of the tenth urite with six claws. \?. Mandibular
appendage with nine to eleven ventral ridges. . . . . . . C. albulus
4. Fore tibia with nine teeth in the dorsal comb. Pedicel of the antenna about
half as long as the scape. Cerci of the tenth urite with four claws. 9. Man-
dibular appendage with nine ventral ridges. . . . . . . €. humatus
92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
THE HOST PREFERENCES OF THE AGAONIDAE1)
“Supremus Genitor Ficui propriam
assignavit cupidinem’’
(HEGARDT, 1749, p. 41)
SPECIFICITY OF THE RELATION BETWEEN FIGS AND FIG WASPS
Theories on the evolution of the symbiosis between figs and fig wasps are
necessarily highly speculative. Fossil data do not provide any clue to the problem,
although a fossil fig wasp (Tetrapus mayri Brues, 1910, p. 16) was described from
the Tertiary (Miocene)?) of Florissant, Colorado, and the presence of fossil fig
leaves is known from the Cretaceous.
Presumably, the fig wasps have descended from gall-forming Chalcidoidea, or
from parasitic Chalcidoidea living on other insects in the flowers or seeds of the
pre-Ficus. The assumption that the interaction of figs and wasps secondarily
became synagonistic — gradually acquiring mutual value and evolutionary signif-
icance for both participants (LEPPIK, 1957, p. 479) — is not new. Although
MULLER (1886b, p. 63) already stated: “die Feigen (haben) einen ursprünglich
einfach ihre Früchte zerstörenden Schmarotzer für thre Kreuzung sich immer
volkommener dienstbar gemacht”, all subsequent theoretical considerations deal
almost exclusively with morphological features of the fig wasps (GRANDI, many
papers, see 1955c; english summary in 1961). MULLER, for that matter, did not
elaborate on his thesis either. Moreover, he was of the opinion that also parasitic
and inquiline wasps could pollinate the fig flowers, especially those of isolated
trees (MULLER, 1883, p. 312). The flight capacity of Blastophaga, and con-
sequently its dispersal, was supposed to be very weak.
Probably, only the Agaonidae serve as pollinators, every species to its peculiar
species of Ficus. The dispersal of the fig wasps does not seem to be such a problem
as might be presumed: PEMBERTON (1934, p. 379) stated that both Pleistodontes
imperialis Saunders and P. froggatti Mayr reached Kauai from Mahu (Hawaiian
Islands) without human assistance). The width of the Kauai Channel is about
one hundred kilometers!
As incidentally remarked above, the relation between figs and fig wasps is
supposed to be highly specific, and it seems to represent a strict one-to-one
relationship. Some evidence of this supposition may be found in the systematic
part of this paper. More arguments, and reputed contra-evidence, objections and
related considerations are discussed below.
Some authors (VAN DER PIJL, 1960, p. 415; BAKER, 1961, p. 378) pointed out
the evolutionary dangers with which every narrow specialist must cope. VAN DER
1) “Agaonidae” and “fig wasps’ are used here in the restricted sense (WIEBES, 1961b),
without including the Sycophaginae, and thus coincide with Agaoninae auctt., and Agaonini
auctt.
2) Lower to Middle Oligocene, according to MAC GINITIE as cited by AXELROD (1954).
3) A similar phenomenon was recorded by CoRNER (1958, p. 18) for the wasps from Ficus
fistulosa Reinw. and F. grossularioides Burm.
J. T. Wreses: Indo-Australian Ceratosolen 93
PiJL is surprised to find the genus Ficus, each species with its own gall wasp, still
flourishing; BAKER, however, seems to change his surprise to disbelief and does
not accept the specificity until more data are available (BAKER, l.c., and in litt.).
Yet, I think that the evidences in favour of a specific relation are fairly sound as
far as any extrapolation in biology goes. Moreover, in my opinion, the “evolution-
ary dangers” are not correctly understood. Admittedly a narrow specialism may
endanger evolution to the extent of making the development of new major types
well nigh impossible, but there is no reason why it should hamper evolution at the
specific level. The great variety of related forms within the genus Ficzs, which
all are bound to the same mode of pollination, and the relatively uniform family
Agaonidae, all species of which live and must develop in the gall receptacles of
Ficus; in short: many species without any clear-cut generic intersections, are what
should be expected as the result of a narrowly specialized and obligate symbiosis,
where other features, as fecundity and dispersal, do not retard evolution.
The experiments on the cross pollination of Ficus carica Linn. and F. pumila
Linn., which showed the perfect compatibility of these species — and which led
BAKER (l.c., p. 379) to the suggestion that many hybrids might occur that are
not recognized as such — are not conclusive. These experiments only show that the
“choice” of the pollinating agent may be the only isolating mechanism between
the fig species (which in this case, however, do not even occur sympatrically).
WILLIAMS (1928, pp. 9, 10) induced Eupristina bakeri Grandi, the wasp from
Ficus forstenii Miq., to crawl into the small figs of F. refusa Linn. (= F. micro-
carpa Linn.f.), the normal inhabitant of which in the Philippines is E. verticillata
Waterston:1)
“A F. retusa fruit was placed in a test tube containing some of the wasp pollina-
tors belonging to F. forstenii, and while these wasps were not constantly watched,
soon a pair of wings was observed protruding from the F. retusa ostiole, where
the wasp's body had not altogether disappeared within, the ovipositor being flat-
tened into some drying liquid discharged from the abdomen. This comparatively
immense E. bakeri wasp finally worked her way within. She did nothing in the
receptacle save walk about prying here and there with her head, using the now
stubby antennae as a wedge apparently in an endeavor to force apart the flowers.
Another similar experiment was unsuccessful.”
Similar experiments were also done with Ceratosolen notus (Baker) — from F.
nota (Blanco) Merrill — in F. barnesii Merrill (= F. botryocarpa Miq.), the
normal pollinator of which is C. corneri Wiebes:
. several wasps had succeeded in gaining complete entrance and stood free
on the inner scales, their abdomen compressed and with sharp lateral angles greatly
suggesting an expulsion of body fluid, and the ovipositor still projecting needle-
like behind. Now to be in readiness for oviposition an effort was made to pull
the ovipositor from its sheath so that it points downwards beneath the forepart of
the abdomen; in this some are immediately successful, curving the abdomen, jab-
bing the ovipositor into the funnel-like stigma of a gall flower and then in
1) I give these and other citations at some length, because they describe the only exper-
iments of this sort; experiments worth while to repeat with more species.
94 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
bringing the abdomen back to the natural, more or less horizontal position the
ovipositor is swung forward out of its sheath; in other cases the wasp has great
difficulty or even fails — possible due to abnormal conditions — in unsheathing
her egg-laying apparatus, bending down the tip of her abdomen and endeavor-
ing to hook her claws into the ovipositor and draw it out.”
and further:
“it seemed that very few ovipositions (?) occurred, and such as they were —
for the conditions seemed unsuitable — occupied in one case about a minute.”
WILLIAMS stated that his experiments showed that it is possible to induce fig
wasps to enter other than their own particular species of fig, although this would
be expected to take place but rarely in nature. In view of the large number of
species of wild figs that may occur in a single habitat, it is, according to WILLIAMS,
not unreasonable to suppose that hybridization might have taken place there.
In addition to this conclusion I may remark that in the experiments the figs were
presented in closer proximity to the wasps than I presume any wasp — with the
possible exception of the inhabitants of strangler figs and their victims — ever
to come near a “foreign” fig receptacle. Probably, the wasps are attracted by the
scent of their own Ficus; this scent may be produced by glands on young leaves
and petioles1).
A questionable point to some authors is whether the wasps carry enough pollen
from a ripe to a young fig for the development of the thousands of ovaries that
may be present in one receptacle, and whether or not the pollen is used at all by
the fig. In a nearly mature female receptacle of F. roxburghii Wall. (= F. auri-
culata Lour.), with general enlargement of the ovaries, CUNNINGHAM (1889, p.
42) found three insects with only one or two shrunken pollen-grains adhering to
them. Contrarily, PEMBERTON (1921, p. 306) stated for Pleistodontes froggatti
Mayr in F. macrophylla Desf. that sufficient pollen is carried over to secure ample
pollination for a great many flowers in each fig. TREUB (1902, p. 137) found the
pollen grains in the female receptacles in F. hirta Vahl: “assez nombreux pour
effectuer la fécondation, sinon de toutes, en tout cas de la grande majorité de
fleurs’. BAKER (1913, p. 68) described the habits of the female Ceratosolen
notus (Baker) in the young fertile figs of F. nota (Blanco) Merr. as: “distributing
to the stigmas the pollen with which her body was thoroughly dusted”.
CUNNINGHAM concluded from his observations that the seeds develop by ad-
ventitious embryony from the nucellus. This agamospermic reproduction needed
the action of the female fig wasp, but it was independent of pollination2). Similar
observations were made by TREUB. He concluded from his cytological investigations
that F. hirta reproduces parthenogenetically. The embryogenesis presumably needed
the action of the insect as a stimulus, but the embryo started its development
without amphimixis.
1) In this, as in the previous part of this paper, I draw freely from data on Ficus given
by CORNER (in litt. to VAN DER VECHT and WIEBES, and personal communication).
2) Serious criticism on the botanical data in CUNNINGHAM’s paper was published by
VAN DER PijL (1934, pp. 179—181). CUNNINGHAM’s remarks on the identity of the wasp
species too, as well as its identification by Woop-MASON with a species of “Euprisizs’’,
are to be regarded with extreme caution.
J. T. WieBEs: Indo-Australian Ceratosolen 95
BUSCALIONI & GRANDI (1938, p. 243) recorded that in F. carica Linn.: “la
cellula ova non fecondato va a male”.
JOHRI & KONAR (1938) contributed a short preliminary note on the embryology
of F. religiosa Linn.
There are many more papers on this problem, an account of which may be
found in the bibliography in JOSEPH (1958), mainly on normal and abnormal
development in the edible fig, but new experiments on wild figs are badly wanted.
Admittedly apogamy and parthenocarpy occur, but I doubt whether these phe-
nomena are of general occurrence under natural conditions.
However this may be — whether the insects give the stimulus by stinging the
stigmata, or bring the stimulus in the form of pollen, or both — there are many
records of figs cultivated in foreign countries that did not fructify until their par-
ticular species of wasps were introduced (F. carica Linn. and Blastophaga psenes
(L.) in California: HOWARD (1901); F. macrophylla Desf. and Pleistodontes
froggatti Mayr in the Hawaiian Islands: PEMBERTON (1921), SWEZEY (1923),
TIMBERLAKE (1924); F. rubiginosa Desf. and P. imperialis Saunders in the Ha-
waiian Islands: SWEZEY (l.c.), TIMBERLAKE (l.c.); etc.1)).
There are some records of one species of Ficus harbouring different species of
Agaonidae, and of one species of fig wasp inhabiting different species of Ficus.
Although it is rather premature to discuss all these cases before the various
genera have been revised, I give an annotated list of these records. Some were
already mentioned by other authors, e.g., NIKOLSKAJA (1956, p. 573), and they
tend to be discussed over and over again in literature. Oddly enough, however,
these reputed cases do not appear to be very important. It is easily understandable
that one species of Ficus can harbour two (sub-)species of Agaonidae in different
parts of its area, or that the host of one species of fig wasp may show some
geographic variation.
Some forms, both botanical and entomological, that are treated as separate spe-
cies, are perhaps only subspecifically distinct. As already stated on p. 85, I treat
the wasps as full species until more material from more localities gives evidence
to the contrary. As to the species of Ficus, some of the forms, although inseparable
in flower characters, are so distinct in the field that, for the time being, CORNER
maintains them as separate species.
By presenting the following list I hope to attain that the attention in future be
concentrated on the important records, and that all records that have no actual
interest to our problem be expelled from the discussion.
1. The record of Blastophaga brasiliensis Mayr from several species of figs, is
based on the following passage from MULLER (1886a, p. 57): “Hier [in Itajahy,
Brazil wurde ein und dieselbe Blastophaga (BI. brasiliensis) in sieben verschiede-
nen Feigen (Ficus II, III, IV, V2), VI, VII, IX) gefunden; sind auch drei dersel-
ben (II, IV, VI) so ähnlich, dass sie vielleicht zur selben Art gerechnet werden
1) The statement by JuDD (1921) on the introduction of fig wasps to the Hawaiian Is-
lands may be of some historical interest.
2) According to MAYR (1906, p. 185), MULLER’s Ficus V is F. dolaria. Ficus dolaria
Martinus, nomen nudum, = F. gomelleira K. & B.
96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
können, so bleiben doch mindestens fünf scharf geschiedene und zum Teil weit
verschiedene, am gleichem Orte wachsende Feigenarten auf eine einzige Blasto-
phaga als wichtigsten!) Bestaübungsvermittler angewiesen”. This statement wants
confirmation.
2. GRANDI (1927a, p. 326) already discussed the case of Blastophaga browni
(Ashmead), recorded from Ficus ulmifolia Lamk. and F. heterophylla Linn., both
from the Philippine Isles. The latter species of Ficus, however, does not occur in
the Philippines, and all Philippine specimens identified with F. heterophylla
actually belong to F. wlmzfolia.
3. JOSEPH (1953c, p. 267) described Blastophaga constabularis Joseph from
Ficus infectoria Roxb. (India, = F. virens Ait.), but from the Sumatran F. 77-
fectoria, B. coronata Grandi (1928b, p. 75) was already known. Judging from
the descriptions, the two are synonyms.
4. The record of Blastophaga grossorum Gravenhorst (= B. psenes (L.)) from
Ficus serrata Forsk. (=F. exasperata Vahl) by Mayr (1885, pp. 153, 179,
“Wüste am rothen Meere”) may well have resulted from the misidentification of
the Ficus. It might have been F. palmata Forsk., which is close to F. carica Linn.,
and which harbours B. vaidi Joseph (1954, p. 401; India), which is close to B.
psenes. MAYR (l.c.) also recorded B. grossorum from F. pseudocarica Miq. (Abys-
sinia) and F. persica Boiss. (Iran), which both are synonyms of F. carica Linn.
5. Blastophaga longicornis Grandi (1926, p. 354) was described from Ficus
rostrata Lamk. (Sumatra, E. JACOBSON, no. 2130), B. dubia Grandi (1926, p.
356) from F. rostrata urophylla Wall. (Java, Hort. Bog., n.VII, G. 46). The
identification of E. J. 2130 with F. rostrata Lamk. (= F. sinwata Thunb.) is cor-
rect; Hort. Bog. n.VII, G. 46 probably is F. rophylla Wall., the correct name
of which is F. heteropleura BI.
6. GRANDI (1916b, p. 225) described the females of Eupristina saundersi
Grandi from Ficus religiosa Linn. (India), from which MAYR (1885, p. 176)
described Blastophaga quadraticeps Mayr (Malaya). GRANDI (1923a, p. 296),
JOSEPH (1953c, p. 277), and JOHRI & KONAR (1955, p. 384) recorded B. qua-
draticeps from Ficus religiosa in India and Ceylon, and I find the same symbiosis
in a sample from Israel (probably introduced).
There is some more confusion as to the host of Eupristina saundersi: the males
were recorded from F. retusa Linn. var. nitida King?) ! GRANDI's host record of
E. saundersi is probably incorrect, as is, most probably, his presumption (1952c,
p. 96) that the host of Pleistodontes semiruficeps Girault (1929, p. 318: “on
Banyan figs, Lord Howe Island”) might be the same as F. religiosa. It might more
likely have been F. macrophylla Desf., or one of its relatives.
7. I find Ceratosolen constrictus (Mayr)3) in Ficus fistulosa Reinw. (Java),
and in F. dimorpha King (Sumatra). The latter, however, is so close to the former,
1) MULLER (see also 1887b, p. 163) considered the possibility that some Idarninae
transmit pollen with the pubescent ovipositor.
2) F. retusa, as used by botanists, is F. microcarpa Linn.f., and it is not genuine F. retusa
Linn., which has been known as F. truncata Miq.
3) For the records of the species of Ceratosolen, I refer to the data given in the system-
atic part of this paper.
J. T. WieBES: Indo-Australian Ceratosolen 97
that it is probably a variety only, but it is insufficiently known botanically.
I have no check on the record of C. hewitti Waterston from F. chartacea Wall.
(Malaya). In the Malayan F. fistulosa, C. hewitti is found, and from F. chartacea
I possess an undescribed species of Blastophaga. Here is a case of a geographic
variation in the wasps (C. constrictus and C. hewitti), that is not, or less, evident
in the Freus.
8. Ceratosolen emarginatus Mayr is found in both Ficus auriculata Lour. and
F. oligodon Mig. The two forms of Ficus are very close. Their ranges seem to
be the same for the greater part, but F. ol/godon extends further south in Malaya,
and hybrids are known from Indo-China, where the forms intergrade. Here is a
case of a variation in the Ficus, which is not evident in the wasp.
The record of C.? crassttarsus: Grandi from Malayan F. roxburghii Wall. (=
F. auriculata Lour.) is probably incorrect.
9. Mayr described Ceratosolen appendiculatus (Mayr) from Ficus umbellata
Hort. Bog., nec Vahl, and C. striatus Mayr from F. variegata BI., both collected
in Java. SOLMS (in MAYR, 1885, p. 193) considered F. umbellata Hort. Bog. to
be the same as F. hirta Vahl, from which, however, Blastophaga javana Mayr is
known (from var. sefosa King; Java).
One specimen in the Leiden Herbarium (sheet no. 908, 188—1895), labelled
“Ficus umbellata Vahl, Kondang, Cult. in Hort. Bog.”, was identified by CORNER
with F. variegata Bl, which supports the suggested synonymy of C. striatus and
C. appendiculatus.
The other figs, recorded to harbour C. appendiculatus, are very close to F.
variegata: F. garciae Elmer and F. sycomoroides Miq. were recently recognized as
varieties, and F. virzdicarpa Corner should perhaps be regarded as a variety too.
The latter is an example of a form that is so distinct in the field, although very
close in structural characters to F. varzegata, that CORNER prefers the two to be
maintained as separate species.
10. Ficus ribes Reinw. (Java) is known to be the host of Ceratosolen crassitarsus
(Mayr). The Philippine variety of F. ribes, var. cuneata (Miq.) Corner, harbours
C. gracilis Wiebes. Here again a geographic variation in casu in both the Ficus
and the wasp.
11. Ceratosolen notus (Baker) is recorded from Ficus nota (Blanco) Merrill and
from F. congesta Roxb., both occurring in the Philippines. F. congesta seems to be
more sylvan in habit than F. rota, but the two are so close that they can only with
difficulty be distinguished in the field and in most cases not at all in herbarium
specimens. The Ficus nota complex is in need of more examination botanically.
12. Ficus hispida Linn.f. harbours C. s. solmsi (Mayr) in Malaya and the
Eastern Archipelago, and on the Asiatic mainland it is inhabited by C. s. marchali
Mayr.
Mayr recorded C. solmsi also from F. canescens Kurz (?, unknown to Cor-
NER, possibly a manuscript name), and Covellia didyma Miq., both from Java
The latter is a synonym of F. lepicarpa BI., which was (see also no. 14) obvious-
ly misidentified by Sous.
13. I cannot find any constant differences between the wasps from Ficus bec-
cari King var. latifolia Corner and those from F. subterranea Corner, and I refer
98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
. both to Ceratosolen humatus Wiebes. The figs, although very close, are distinctly
different in leaf-shape, and the distinction is approximately of the same magnitude
as that between F. beccarii latifolia and F. uncinata Becc. var. strigosa Corner,
which harbours a distinct form of Ceratosolen, C. albulus Wiebes.
This problem needs further study.
14. Ceratosolen bisulcatus (Mayr) was described from Ficus lepicarpa BI.
(Java). In the receptacles of this Ficus, however, I find C. vechti Wiebes (Java,
Sumatra). C. bisulcatus inhabits the figs of F. septica Burm.f. in Java, Bali, and
New Guinea, whereas in the Philippine F. septica, C. jucundus Grandi is found.
SOLMS, who identified Mayr’s Ficus, evidently did not understand F. lepicarpa
properly, as is, moreover, seen by his identification of the host of C. solmsi
(see no. 12) with Covellia didyma (= F. lepicarpa).
15. Ceratosolen fusciceps (Mayr), known to be the symbiont of Ficus racemosa
Linn. (from India to Queensland), was also recorded from F. /ucescens Miq.
(=F. racemosa var. elongata (King) Barrett) and F. covellii Roxb. (not known
to CORNER).
From Australian F. glomerata (= F. racemosa), GIRAULT described Blasto-
phaga nigriscapus Girault (1925, p. {2}) and B. niveipes Girault. Thanks to the
kindness of Dr. E. F. Riek, Canberra, I had the opportunity to study some female
specimens of B. niveipes, which might actually belong to C. fusciceps. I did not
see B. nigriscapus. Material from Queensland, ex Ficus racemosa (coll. WILLIAMS,
HSPA), is identical with C. fusciceps; indeed, this species seems to be little varia-
ble over the whole of its very large range.
Ceratosolen mysorensis Joseph (India) is, in my opinion, identical with C.
fusciceps (Mayr). The species was recorded from F. mysorensis Hayne (= F.
drupacea Thunb. var. pubescens (Roth) Corner), but, a year later, Eupristina
belgaumensis Joseph (1954, p. 409) was described from the same Ficus. As
F. drupacea belongs to the group of strangler figs allied to F. benghalensis Linn.,
which all seem to harbour species of Eupristina, I presume the record of E.
belgaumensis to be correct. F. drupacea may have been growing on a tree of F.
racemosa, thus resulting in the confusion!) of the Ficus species.
16. SAUNDERS (1883, p. 6) described Eupristina masoni Saunders from Ficus
indica Linn. (India). In 1928 (a, p. 81), GRANDI recorded E. ?masoni from
F. benghalensis Linn., which, however, he considered to be the same as F. indica.
From F. benghalensis was recorded (JOSEPH, 1953c, p. 282) E. grassii Grandi, of
which the author stated (GRANDI, 1928c, p. 225): “è specie affine e, probabil-
mente, identica a Masoni Saunders”.
In this, as in the following two problems, one must be aware of the fact that
the host relations of the genus Ezpristina can only be solved by a close cooperation
between the botanist and the entomologist. The host Ficzs are very close, and
were not infrequently confused. The same is true for the species of Eupristina.
17. Ficus benjamina Linn. (Sumatra) is known to harbour Eupristina jacobsoni
1) Either by the collector, in taking wasps from the one species, and botanical material
from the other; or by the wasp, which might be unable to distinguish between fig receptacles
of the different species, once the tree is found (?, cf. the experiments by WILLIAMS,
discussed in this paper, p. 93).
J. T. WieBEs: Indo-Australian Ceratosolen 99
Grandi (1926, p. 358), which is very close to E. koningsbergeri Grandi from
F. benjamina var. comosa King (GRANDI, 1917, p. 52; Java). This variety is
distinguished by having a much larger, rounded fig than typical F. benjamina.
JOSEPH (1954, p. 415) recorded E. koningsbergeri from F. benjamina in India.
18. A similar problem exists in the wasp species recorded from Ficus retusa
Linn. (= F. microcarpa Linn.f., see note on p. 96): Eupristina verticillata
Waterston is known (GRANDI, 1926, p. 358) from the typical form (Sumatra,
and presumably, sub Grandiella (WILLIAMS, 1928, p. 13), also from the Philip-
pines), whereas IsHI (1934, p. 85) described E. okinavensis Ishii from the Ja-
panese var. uitida Thunb. (see also this list, no. 6).
19. In 1961 (a, p. 231) I described Agaon paradoxum modestum Wiebes from
Ficus brachypoda Hutch. (Uganda). The nominate subspecies is known (GRANDI,
1952a, p. 31) from F. ovata Vahl (West Africa). The figs, however, are so
close that CORNER would not insist on the specific diversity of F. ovata and F.
brachypoda.
20. For the sake of completeness I mention the cases of Pleistodontes spp.,
which were recorded from Australian figs (see GRANDI’s catalogue, 1955b, p.
129). The confusion between Ficus columnaris Müll.1), F. macrophylla Desf.,
F. australis Willd.2), F. rubiginosa Desf.3), and F. eugenioides Müll.4) (PEM-
BERTON, 1944, p. 17), recorded by various authors as hosts of Pleistodontes frog-
gatti Mayr, P. imperialis Saunders, and P. nigricaput Girault, may be due to a
misunderstanding of the taxonomic status of the Ficus, and also to some mis-
interpretation of the wasp species.
PHYLOGENETIC SPECIFICITY
Additional, indirect evidence in favour of a specific relation between fig wasps
and figs is supplied by the observation that related species of figs harbour related
species of fig wasps. Mutatis mutandis, this principle was expressed in “FAHREN-
HOLZ's Rule” of parasitologists, in which is stated (I quote from SZIDAT, 1956,
p- 243), that “bei stetigen Parasiten aus der Systematik der Parasiten meist un-
mittelbar auch auf die Verwandtschaft der Wirte schliessen {lässt}.
Tables II and III give a survey of the data on the host records of the Agaonid
genera5), and of most species of Ceratosolen, respectively.
All host records of the Agaonidae known to me are incorporated in Table II.
Specific records of the African fig wasps were mentioned by WIEBES (1961b,
table I); specific records of the American Agaonidae will be listed in a later paper.
1) = Ficus macrophylla Desf.
2) = Ficus rubiginosa Desf.
3) Ceratosolen silvestrianus Grandi (1916a, p. 160) was recorded from F. ferruginea
Desf. (Africa!), which is the same as the Australian F. rubiginosa Desf. (with Pleistodontes
imperialis Saunders).
4) = Ficus obliqua Forstf. The sample, from which Blastophaga greenwoodi Grandi
(1931, p. 8) was recorded, may have been F. prolixa Forst.f.
5) Host records are known of species of the following genera or subgenera: Blastophaga
Gravenhorst, Elisabethiella Grandi, Waterstoniella Grandi, Julianella Grandi, Valentinella
Grandi, Kradibiella Girault, Ceratosolen Mayr, Eupristina Saunders, Tetrapus Mayr, Allo-
triozoon Grandi, Agaon Dalman, and Pleistodontes Saunders.
100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
As to the Indo-Australian wasps I have taken into account many species in our
collection that are as yet only generically identified.
Many records had to be incorporated on which I have no check. In the evaluation
of the data from the tables another reservation must be borne in mind. The
groups of Ficus and the genera or species of the Agaonidae are presented in a
linear sequence, and one cannot ever hope to depict but poorly the intricate multi-
dimensional relationships in a linear system.
A similar, more conclusive list of the host records should be given when more
genera are monographically revised, and when the classifications of Ficus and
the Agaonidae have been reconsidered in a joint revaluation by the botanist and
the entomologist.
The following notes may accompany this first, tentative draft of the host
records of the Agaonidae.
The subgenera Urostigma and Pharmacosycea; tentative remarks on the African,
American, and Australian genera of the Agaonidae
1. The relations within the subgenus Urostigma, as well as those in the Agaon-
idae, are very intricate, and they cannot be evaluated properly in this cursory re-
view. I want to state here, that I am not satisfied with the usual generic division of
the Agaonidae. In my opinion, Elisabethiella should not be regarded as a sub-
genus of Blastophaga sl; Eupristina, on the other hand, probably should; it
appears to be the Asiatic analogon of American Julianella.
2. The record of Ceratosolen silvestrianus Grandi from the section Malvanthera
(Ficus rubiginosa Desf.; Africa, see WIEBES, 1961a, p. 239) is doubtful. The
record of a species of Ceratosolen from the section Galoglychia is based on the
description of C. myersi Grandi (1955a, p. 92) from F. rhodesiaca Warb.
(Africa). All other African species of Ceratosolen live in figs of the subgenus
Sycomorus.
3. Some species of the section Urostigma and some of the subsection Varinga,
subgenus Ficus, are known to occur in Africa, but the pollinating wasps are not
known. The Indo-Malayan species harbour Blastophaga s.str.
4. Two species of Elisabethiella were recorded from figs of the section Galo-
glychia. One species (Blastophaga (Elisabethiella) wanei Risbec, 1951, p. 386)
was recorded from Ficus gnaphalocarpa Steud. (subgenus Sycomorus), but I have
reasons to assume this record to be incorrect.
5. The section Oreosycea harbours Blastophaga, but from one species (Ficus
pritchardii Seem., of uncertain position) a species of Ceratosolen is known (p.
8), and from F. vasculosa Wall. ex Mig. (Singapore, ? cultivated) I have an
insect that seems to belong to Pleistodontes.
6. The record of Tetrapus is based on the observation by MULLER (1887a, p.
X) that Tetrapus appears to be limited to Pharmacosycea, but no specific records
are available.
The subgenera Ficus and Sycomorus, and the Agaonid genera Blastophaga,
Kradibiella, and Ceratosolen
7. As far as known, all series of the section Ficus harbour Blastophaga s.str.,
J. T. WieBEs: Indo-Australian Ceratosolen 101
TABLE U. Ficus AND THE GENERA OF THE AGAONIDAE
Ficus Linn. Agaonidae
Urostigma (Gasp.) Miq.
Urostigma Blastophaga
Leucogyne Corner Eupristina
Conosycea (Miq.) Corner Blastophaga, Eupristina, Waterstoniella
Stilpnophyllum Endl. Blastophaga
Malvanthera Corner Pleistodontes, Ceratosolen
Galoglychia (Gasp.) Endl. Elisabethiella, Allotriozoon, Agaon,
Ceratosolen
Americana Miq. Julianella, Valentinella
Pharmacosycea Miq.
Pharmacosycea Tetrapus
Oreosycea (Miq.) Corner Blastophaga, Ceratosolen, Pleistodontes
Ficus
Ficus
Pseudopalmeae Corner Ceratosolen
other series Blastophaga
Sycidium Miq.
Sycidium
Prostratae Corner Ceratosolen
Pungentes Corner Ceratosolen
Phaeopilosae Corner Blastophaga
Copiosae Corner Blastophaga
Scabrae Miq. Blastophaga, Kradibiella
Varinga (Miq.) Corner Blastophaga
Palaeomorphe (King) Corner Blastophaga
Rhizocladus Endl. Blastophaga
Kalosyce (Miq.) Corner Blastophaga
Sinosycidium Corner
Adenosperma Corner
Neomorphe King Ceratosolen
Sycocarpus Miq. Ceratosolen
Sycomorus Miq. Ceratosolen
with the exception of the Psendopalmeae: Ceratosolen bakeri Grandi lives in the
figs of Ficus pseudopalma Blanco. This wasp has aberrant characters for the genus
Ceratosolen, but it appears to be related to C. abnormis and to the C. armipes
group. The Ficus was classified in the section Ficus because of its bistaminate male
flowers, but it does show some relationships with F. dammaropsis Diels, the host
of C. abnormis Wiebes.
8. The species of Blastophaga from the various series of the sections Ficzs,
Sycidium, Rhizocladus, and Kalosyce belong to several species groups. The general
impression is, that there is a good correspondence between the classifications of
the figs and the fig wasps, but a more positive statement cannot be given before
the species of Blastophaga have been revised.
9. A species of Kradibiella was recorded from the Australian Ficus stephano-
carpa Warb. (= F. coronata Spin.) by WAKEFIELD (1960), and its description
will shortly be given by Riek. I should add here, that I am not convinced of the
need for a distinct genus Kradzbiella: it would seem to represent merely a species
group in the genus Blastophaga. The alternative to the inclusion of Kradzbiella in
Blastophaga is to recognize probably as many as five or six other genera.
102 ‘TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
10. The greater part of the Indo-Malayan and Papuan species of Ceratosolen
live in figs of the sections Neomorphe and Sycocarpus, but some are known from
two series of Sycidium. It is interesting to note that these series have usually been
placed in Sycocarpus, and only recently have been assigned to Sycidium. Botanic-
ally, these groups “point to an ancestor which would combine Sycidium with
Sycocarpus and Sycomorus {here including Neomorphe |} (CORNER, 1958, p. 31).
In my opinion, the wasp from the series Prostratae (Ceratosolen gravelyi
Grandi, ex Ficus semicordata Ham. ex Smith) connects those from the section
Neomorphe with those from the subgenus Sycomorus (see Table III), and the
wasps from the series Pungentes (C. pygmaeus Grandi, and C. nanus Wiebes, from
F. minahassae Mig. and F. pungens Reinw., respectively) appear to be related to
the wasps from the section Sycocarpus. The botanical parallel is not clear.
11. In figs of the series Phaeopilosae and Copiosae, I find the grades between
Blastophaga and Ceratosolen alluded to in a previous part of the present paper
(p. 4). These are the groups that should be studied before the genera Cerato-
solen and Blastophaga can be more exactly defined.
TABLE ul. Ficus (Neomorphe, Sycocarpus, AND Sycomorus) AND THE INDO-MALAYAN AND
PAPUAN SPECIES OF Ceratosolen
Ficus Linn. Ceratosolea Mayr
C. appendiculatus group
subgenus Sycomorus Mig 1)
F. racemosa Linn. C. fusciceps (Mayr)
do., var. elongata (King) Barrett C. fusciceps (Mayr)
(subgenus Ficus)
sectio Neomorphe King
series Auriculatae Corner
F. auriculata Lour. C. emarginatus Mayr
F. oligodon Miq. C. emarginatus Mayr
series Variegatae Corner
subseries Variegatae
F. variegata BI. C. appendiculatus (Mayr)
do., var. garciae (Elmer) Corner C. appendiculatus (Mayr)
do., var. sycomoroides (Miq.) Corner C. appendiculatus (Mayr)
F. viridicarpa Corner C. appendiculatus (Mayr)
subseries Laciniatae Corner
F. semivestita Corner C. grandii Wiebes
sectio Sycocarpus Miq.
C. armipes group
subsectio Auriculisperma Corner
series Cynaroides Corner
F. cynaroides Corner C. sordidus Wiebes
series Theophrastoides Corner
series Vitienses Corner
F. salomonensis Rech. C. boschmai Wiebes
subsectio Papuasyce Corner
F. microdictya Diels C. spec.
F. itoana Diels C. armipes Wiebes
1) For convenience’ sake, I begin this table with the subgenus Sycomorus, although it is
usually placed at the end of the botanical system (cf. Table II).
J. T. WieBEs: Indo-Australian Ceratosolen
Ficus Linn.
subsectio Dammaropsis (Warb.) Corner!)
F. dammaropsis Diels
subsectio Lepidotus Corner
subsectio Macrostyla Corner
subsectio Sycocarpus
series Longetuberculatae Sata
F. ribes Reinw. ex Bl.
do., var. cuneata (Miq.) Corner
F. botryocarpa Miq.
do., var. subalbidoramea (Elmer) Corner
series Tuberculifasciculatae Sata
subseries Praestantes Corner
F. praestans Corner
subseries Calopilinae Corner
F. calopilina Diels
F. bernaysii King
F. iodotricha Diels
F. obpyramidata King
subseries Congestae Corner
F. congesta Roxb.
F. nota (Blanco) Merrill
F. subcongesta Corner
do., var. symmetrica Corner
subseries Hispidae Corner
F. hispida Linn.
F. moderata Corner
F. hispidioides S. Moore var. succosa Corner
subseries Axillares Corner
F. lepicarpa BI.
subseries Fulvidulae Corner
F. cereicarpa Corner
F. francisci Winkler
F. treubii King
subseries Geocarpicae Corner
F. uncinata Becc. var. strigosa Corner
F. beccarii King var. latifolia Corner
F. subterranea Corner
F. stolonifera King
subseries Tuberculifasciculatae
F. fistulosa Reinw.
do.
do., var. tengerensis O.K.
F. dimorpha King
F. septica Burm.f.
do.
103
Ceratosolen Mayr
C. abnormis group
C. abnormis Wiebes
C. crassitarsus group, p.p.
. crassitarsus (Mayr)
. gracilis Wiebes
. corneri Wiebes
. corneri Wiebes
VaAAO
. praestans Wiebes
. calopilinae Wiebes
. hooglandi Wiebes
. iodotrichae Wiebes
. nugatorius Grandi
. notus (Baker)
. notus (Baker)
. orientalis Wiebes
. ?orientalis Wiebes
MOA Ada ©
C. solmsi group, p.p.
C. solmsi (Mayr)
(C. crassitarsus group, continued)
C. moderatus Wiebes
C. dentifer Wiebes
C. vechti Wiebes
C. pilipes group, p.p.
C. pilipes Wiebes
C. josephi Wiebes
(C. solmsi group, continued)
C. brongersmai Wiebes
(C. pilipes group, continued)
C. albulus Wiebes
C. humatus Wiebes
C. humatus Wiebes
C. spec. (group?)
C. pygmaeus group, p.p.
C. constrictus (Mayr)
C. hewitti Waterston
C. constrictus (Mayr)
C. constrictus (Mayr)
C. bisulcatus group
C. bisulcatus (Mayr)
C. jucundus Grandi
1) By CORNER (1960, p. 38) placed between subsections Auriculisperma and Pomifera
(now, pp. = Papuasyce, and p.p. = Lepidotus; see CORNER, 1962, pp. 395— 396).
104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
12. The wasps from the sections Neomorphe and Sycocarpus and the subgenus
Sycomorus are listed in Table III.
The species of the Ceratosolen appendiculatus group live in the receptacles of
the section Neomorphe and in those of the subgenus Sycomorus, and one species
is known from the series Prostratae (see no. 10). The occurrence of a group of
so closely related species, as the C. appendiculatus group apparently is, in the figs
of both dioecious Neomorphe and monoecious Sycomorus, would suggest that the
floral characters in which Neomorphe is very close to Sycomorus are more im-
portant than the distribution of the flowers in the receptacles. A parallel is found
in F. microdictya Diels, which is the only monoecious species in dioecious Syco-
carpus.
13. The wasps from the section Sycocarpus can be classified in two large divis-
ions, which coincide with the first three subsections of Sycocarpus (no records
are known from the subsections Lepidotus and Macrostyla), and the subsection
Sycocarpus, respectively.
Some characters of the species of the C. armipes and C. abnormis groups, which
form the first large division, suggest of a relationship with C. bakeri (see no. 7).
I am inclined to regard the common character of the C. armipes group and the
C. pilipes group, the tibial comb of the female fore leg, as the result of a parallel
evolution, and not as an indication of a close relationship.
14. The second large division of Ceratosolen contains the C. crassitarsus, C.
solmsi, C. pilipes, and C. bisulcatus groups.
In this assemblage, the C. pzlipes group is aberrant in the female and (less) in
the male sex, and the C. solmsi and C. bisulcatus groups are distinct in the male
sex only.
In general, the classifications of Sycocarpus and the groups of Ceratosolen run
parallel, be it that the two species of the C. solmsi group break the sequence. C.
solmsi, however, seems to be closest to the species of the C. crassitarsus group,
amongst which it is listed, and C. brongersmai shows some connections with the
C. pilipes group. I will not insist on the value of the C. so/msz group, which had
to be established on the entomological evidence, but may prove to be redundant
when more data are available.
The same reservation must be made for the C. bisulcatus group, which is based
on the bilobate epistomal margin of the males. The subseries Tuberculifasciculatae
contains, next to Ficus septica, the hosts of C. constrictus and C. hewittt, which
I classify with the C. pygmaeus group. The connections with two series of the
subsection Sycidium (see no. 10) and with F. pritchardii (see no. 5) are not clear
botanically. The problem of the relationships between C. constrictus and C. hewitti
on the one hand, and C. bisulcatus and C. jucundus on the other hand, is the more
confusing as in the Philippines F. fistz/osa seems to grade into, or hybridize with,
F. septica.
The figs of the subseries Fulvidulae and Geocarpicae harbour C. brongersmai,
already mentioned above, and the species of the C. pilipes group. The C. pilipes
group consists of two pairs of species, which correspond with the subseries of the
hosts. A fifth species, from F. stolonifera, could not be described on the scanty
and badly preserved material, but it does not show what I assume to be the most
J. T. WIEBES : Indo-Australian Ceratosolen 105
important character of the C. pzlipes group, viz., the blunt tooth in the dorsal
comb of the female fore tibia.
The sequence in which I listed the species of the C. crassitarsus group, and in
which I tried to depict some of the relationships, does not correspond with the
list of hosts as classified by CORNER. Although the correlation might be better in
a multidimensional system, some species of wasps defy the classification suggested
by the botanist. C. corneri, for instance, is so abnormal in the male sex that it is
difficult to accept its classification with C. crassitarsus and C. gracilis, which,
moreover, do not support the close relationship between their hosts.
The wasps of Ficus calopilina, F. nota, and subcongesta are so close that I am
doubtful of their specific diversity, although the figs are classified in two distinct
subseries.
Ceratosolen nugatorius and C. dentifer, finally, both show the peculiar tibial
spur in the male hind leg, but it may be that these spurs are not homologous. I
could accept the classification as suggested in the botanical system.
Tentative remarks on the Agaonidae
15. Omitting all doubtful cases, a general survey would show the species of the
genus Blastophaga as pollinators in three subgenera of Ficus, viz., Urostigma,
Pharmacosycea, and Ficus. In all three, different types have evolved: several sub-
genera of Blastophaga in Asia (Blastophaga, Eupristina, Waterstoniella), Pleis-
todontes in Australia, Julianella, Valentinella, and Tetrapus in America, and
Elisabethiella, Agaon and Allotriozoon in Africal). The parallel evolution of fig
wasps with long faces and denticulate mandibles in the female sex, in Australia
(Pleistodontes), Africa (Agaon), and America (Tetrapus), is mentioned here as
a curious fact, well worth of closer research and comparison.
Ceratosolen appears to consist of three original stocks, roughly coinciding with
Neomorphe and Sycomorus, the first part of the section Sycocarpus, and the
subsection Sycocarpus, respectively. The genus Ceratosolen appears to be a derivate
of Blastophaga, and seems to be still connected with it in the wasps from the sub-
section Syczdium.
CONCLUDING REMARKS
I shortly summarize the line of thought which led to the research on figs and
fig wasps as presented in the preceding chapters.
To find in a taxonomical revision of a group of fig wasps a high specificity in
the close relation between fig wasps and their hosts is to ask the question whether
or not this absolute specificity is of general occurrence in the family. It appears
that most host records confirm to the principle; others do not, but these are
either not documented, and therefore they have no evidential value, or there is
reason to doubt the taxonomical evidence.
The next problem is the nature of the specificity. In the chapter relating to this
matter it was suggested that there exists a phylogenetic specificity, in other words,
Ficus and the Agaonidae are supposed to have evolved together. The evidences in
1) Blastophaga s.str. is not yet known from Africa, except for the introduced Blastophaga
psenes (L.).
106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
|
favour of a phylogenetic specificity are convincing, as seen in the classification of |
the larger groups. The parallelism in the classification of the smaller divisions, and
in that of the species, is less satisfactory. This, however, does not affect the hy-
pothesis of the phylogenetic specificity, as both the botanical and the entomological
classifications should be reconsidered later. For the time being, CORNER and my-
self try to base our classifications of figs and fig wasps, as much as possible,
exclusively on botanical and entomological evidence, respectively. In the long end
we search for a classification that satisfies both the entomologist and the botanist. |
Should such a system be found we may turn the argument and open the discus-
sion on the principles of our classification and on the evolution of the symbiosis.
This is, of course, what we have in mind and what was stated before by VAN DER
VECHT (1956, p. 103) and CORNER (1960, p. 370). Our taxonomic revisions are
intended to be means to that end.
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INDEX
New specific names are indicated by an asterisk *; synonyms in italics
)
AGAONIDAE
*abnormis 25 (figs.), 28 (figs.), 32, 86, appendiculatus(a) 4, 19, 85, 89, 97, 102
[89, 101, 103 *armipes 22 (figs.), 27, 86, 89, 102
Agaon 99, 101, 105 bakeri (Cer.) 28 (figs.), 32, 86, 90, 101
*albulus 75, 79 (figs.), 87, 91, 98, 103 bakeri (Eupr.) 93
Allotriozoon 99, 101, 105 belgaumensis 98
110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
berlandi 68 *josephi 73 (fig.), 74, 76 (figs.), 87, 91,
*bianchii 18 (figs.), 26, 86, 89 [103
bisulcatus(a) 40 (figs.), 81, 87, 90, 98, jucundus 81, 83, 87, 90, 98, 103, 104
[104 Julianella 99, 100, 101, 105
Blastophaga 3, 99, 100, 101, 105 koningsbergeri 99
*boschmai 15 (figs.), 23, 86, 89, 102 Kradibiella 99, 101
brasiliensis 95 longicornis 96
*brongersmai 68, 70 (figs.), 87, 90, 103, marchali 64, 66 (figs.), 67, 68, 87, 97
[104 marshalli 7 (figs.), 8, 85, 88
browni 96 masoni 98
*calopilinae 54, 59, 61 (figs.), 86, 91, 103 mayri 92
Ceratosolen 3, 99, 100, 101, 102, 105 megarhopalus 84
Ceratosolens 4 *moderatus 46 (figs.), 47, 86, 91, 103
Ceratosolensia 4 modestum 99
coeca 3 myersi 100
constabularis 96 mysorensis 17, 98
constrictus(a) 7 (figs.), 12, 85, 89, 96, “nanus 9, 10 (figs.), 85, 88, 102 |
[103, 104 nigricaput 99 |
*corneri 62, 63 (figs.), 86, 90, 93, 103, nigriscapus 98
[105 niveipes 17, 98 |
coronata 96 notandus 19, 20 |
crassitarsus 35, 83, 86, 90, 97, 103, 105 notus 54, 55 (figs.), 86, 90, 93, 94, 97, |
*dentifer 43 (figs.), 44, 86, 90, 103, 105 [103 |
dubia 96 nugatorius 40 (figs.), 42, 86, 90, 103, |
effractarius 16 [105 |
elisabethae 84 okinavensis 99 |
Elisabethiella 99, 100, 101, 105 “orientalis 54, 57, 58 (figs.), 86, 91, 103
emarginatus 16, 85, 89, 97, 102 paradoxum 99 |
Eupristina 99, 100, 101, 105 aes Wil, 75 (Be), Sy, Di, 10%
Eupristis 94 Pleistodontes 99, 101, 105
ficophaga 4, 20 *praestans 49 (figs.), 86, 91, 103
froggatti 92, 94, 95, 99 psenes 95, 96, 105
fusciceps 16, 85, 89, 98, 102 pygmaeus 7 (fig.), 8, 85, 88, 102
fuscipes 16 quadraticeps 96
*gracilis 31 (figs.), 35, 86, 91, 97, 103, saundersi 96
[105 semiruficeps 96
Grandiella 99 silvestrianus 99, 100
*grandii 13 (figs.), 21, 85, 89, 102 solmsi 64, 65, 66 (figs.), 67, 87, 90, 97,
grassii 98 [103, 104
gravelyi 14, 85, 89, 102 *sordidus 25 (figs.), 30, 86, 89, 102
greenwoodi 99 striatus 19, 97
grossorum 96 Sycocrypta 3
hewitti 7 (fig.), 12, 14, 85, 89, 97, 103, Tetrapus 99, 100, 101, 105
[104 vaidi 96
*hooglandi 34 (figs.), 38, 86, 91, 103 Valentinella 99, 101, 105
*humatus 78, 82 (figs.), 87, 91, 98, 103 *vechti 37 (figs.), 39, 84, 86, 91, 98, 103,
imbecillus 14 a il
imperialis 92, 95, 99 verticillata 93, 99
*jodotrichae 51, 52 (figs.), 86, 91, 103 wanei 100
jacobsoni 98 Waterstoniella 99, 101, 105
javana 97
Ficus
acidula 17 Auriculatae 102
Adenosperma 101 Auriculisperma 102
Americana 101 australis 99
auriculata 16, 94, 97, 102 Axillares 103
J. T. WieBEs: Indo-Australian Ceratosolen
barnesit 62, 93
beccarii 78, 97, 103
benghalensis 98
benjamina 98
bernaysii 38, 103
botryocarpa 62, 93, 103
brachypoda 99
calopilina 59, 103, 105
Calopilinae 103
canescens 65, 97
carica 93, 95, 96
cereicarpa 71, 103
chartacea 14, 97
chrysolepis 84
columnaris 99
comosa 99
congesta 54, 97, 103
Congestae 103
Conosycea 101
Copiosae 101, 102
coronata 101
covellii 16, 98
cuneata 35, 97, 103
cunia 14
Cynaroides 102
cynaroides 30, 102
Dammaropsis 103
dammaropsis 32, 101, 103
didyma 65, 97
dimorpha 12, 96, 103
dolaria 95
drupacea 98
ebretioides 20
elongata 17, 98, 102
endothrix 62
eugenioides 99
exasperata 96
ferruginea 99
Ficus 100, 101, 102, 105
fistulosa 12, 14, 92, 96, 103, 104
forstenii 93
francisci 74, 103
Fulvidulae 103, 104
Galoglychia 100, 101
garciae 97, 102
Geocarpicae 103, 104
glomerata 16, 98
gnaphalocarpa 100
gomelleira 95
grossularioides 92
hauili 83
heterophylla 96
heteropleura 96
hirta 94, 97
hispida 64, 97, 103
Hispidae 103
hispidioides 44, 103
indica 98
infectoria 96
integrifolia 20
iodotricha 51, 103
itoana 27, 102
Kalosyce 101
Laciniatae 102
latifolia 78, 97, 103
lepicarpa 39, 81, 97, 98, 103
Lepidotus 103
Leucogyne 101
longipedunculata 84
Longetuberculatae 103
lucescens 16, 98
macrophylla 94, 95, 96, 99
Macrostyla 103
Malvanthera 100, 101
merrilli 35
microcarpa 93, 96, 99
microdictya 83, 102
minahassae 8, 102
moderata 47, 103
mysorensis 17, 98
Neomorphe 101, 102
nitida 96, 99
nota 54, 93, 94, 97, 103,
obliqua 99
obpyramidata 42, 103
obtusa 32
oligodon 16, 97, 102
Oreosycea 100, 101
ovata 99
Palaeomorphe 101
palmata 96
Papuasyce 102, 103
persica 96
Phaeopilosae 101, 102
Pharmacosycea 100, 101
Pomifera 103
praestans 49, 103
Praestantes 103
pritchardii 8, 100
prolixa 99
Prostratae 101, 102
pseudocarica 96
pseudopalma 32, 101
Pseudopalmeae 101
pubescens 98
pumila 93
pungens 9, 102
Pungentes 101, 102
racemosa 16, 98, 102
religiosa 95, 96
retusa 96
retusa 93, 96, 99
, 104
105
2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 1, 1963
Rhizocladus 101
rhodesiaca 101)
ribes 35, 97, 103
rostrata 96
roxburghii 16, 83, 94, 97
rubiginosa 95, 99, 100
salomonensis 23, 102
satterthwaitei 54
Scabrae 101
semicordata 14, 102
semivestita 21, 102
septica 81, 83, 98, 103, 104
serrata 96
setosa 97
Sinosycidium 101
sinuata 96
stephanocarpa 101
Stilpnophyllum 101
stolonifera 84, 103, 104
storcki 8
strigosa 75, 98, 103
subalbidoramea 62, 103
subcongesta 57, 103, 105
subopposita 12 |
subterranea 78, 97, 103
succosa 44, 103
Sycidium 101
Sycocarpus 101, 102, 103, 104
sycomoroides 20, 97, 102
Sycomorus 100, 101, 102, 104
symmetrica 57, 103
tengerensis 13, 103
Theophrastoides 102
treubii 68, 103
truncata 96
Tuberculifasciculatae 103, 104
ulmifolia 96
umbellata 19, 97
umbonata 19
uncinata 75, 98, 103
urophylla 96
Urostigma 100, 101
variegata 20, 97, 102
Variegatae 102
Varinga 100, 101
vasculosa 100
virens 96
viridicarpa 21, 97, 102
Vitienses 102
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G. C. D. GRIFFITHS. — A revision of the Palaearctic species of the nigripes group
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4 i a
A
A REVISION OF THE PALAEARCTIC SPECIES OF THE-
NIGRIPES GROUP OF THE GENUS AGROMYZA FALLEN
(DIPTERA, AGROMYZIDAE)
BY
GEERD IGRIFEMMES NBA ERE:S
88a, Avondale Avenue, East Barnet, Herts., England
CONTENTS
tro et 0 mgee 111
GHALAGtEriSatONGOMMEWCIOUPRT N tete 112
GO PRO 722277 Denm MEIS eN nn 112
Can we 24222 a alléen 113
The nigripes group outside the Palaearctic region ........................................ 114
Comments on earliermtreatmentuotathe! SOUPE TT neater e eee emenc cesse 114
POE ne RE PRE IT OLII re Eee 115
TO stre ao ns Iig eene 117
Kes tor delend HEAO Aer IE Re OI aree iii 119
AUSTERO OR LO RR 119
EA VA CM Eee done 121
ID ESPOSTE I TR NI II ente 122
PATASItE SAM E TAO DE II. BRERA TN IONE gE NM ite IIa 159
INcknowledsementsun a ehe denne TT 164
SU AA TV ES RE te RL ane SR no SU 164
RENES Socks 165
INTRODUCTION
The species of the n7gripes group of the genus Agromyza Fallen are exclusively
feeders on Gramineae, including the cultivated forms. They often come to the
notice of the applied entomologist, and there are numerous references to
them in the literature. This paper, although written primarily from a taxonomical
point of view, is intended also to assist the work of the applied entomologist.
Past work in this field has sometimes been confused through the author failing to
realise that he was dealing with more than one species with different life-histories;
the host-relationships too have not hitherto been very clear. It is hoped that this
paper will provide a sound taxonomic basis for future work.
The need for a complete revision of the group has been apparent for some time.
The external characters of the adult do not provide many good specific differences,
and it was clear that an attempt to use the characters of the male genitalia for
classification needed to be made. These had not been employed by HENDEL
(1931). Also the larval key hitherto in use, originating from DE MEIJERE (1943),
has proved to be almost unworkable. I have decided to make a fresh start with a
new key.
The terminology used for the external morphology of the adult in this paper is
113
114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
an English version of that of HENDEL (1931), as has been commonly used for the
Agromyzidae. The genitalia and larval terminology largely follow Frick (1952),
though I have preferred the term “spine” to FRICK’s “cuticular process”. Figs. 1
and 2 explain the genitalia terminology. The plant names are as given in CLAPHAM,
TUTIN and WARBURG (1952).
CHARACTERISATION OF THE GROUP
The term “nigripes group” is used in this paper in a narrow sense to include
those grass-feeding species of Agromyza in which the costa reaches to m ‚+ 9.
It is closely related to the ambigua group, in which the costa only reaches tr, ‚ 5.
This distinction may ultimately prove untenable on a world basis, but it is a
convenient one for dealing with he Palaearctic species. The distinctive characters
of these two groups may be summarised as follows.
Group of nzgripes Meigen
External morphology
Lunule very narrow, occupying less than 1/4 of the distance from the antennal
base to the front ocellus: ocellar triangle coming to a point well before the front
ocellus. Dorsocentrals basically six (four postsuturals and two presuturals),
decreasing markedly in length forwards: in most species only three or four of the
postsuturals are longer than the acrostichals. Mesonotum more or less shining.
Inner post-alar bristle present. Legs coarsely haired: middle tibiae with a pair of
outstanding posterodorsal bristles. Costa reaching mj + 9.
Male genitalia
Cerci elongate, club-shaped, with long hairs at their tips. Ninth sternite reaching
1/,— 2/3 of the length of the aedeagal apodeme. Postgonites small, rounded or
club-shaped, closely adpressed to the pregonites: pregonites completely fused with
the ninth sternite. Phallophore small, more or less triangular. Basiphallus consisting
of a pair of broad plates, but subject to various modifications. Distiphallus of
characteristic S-shape.
Larva
No patch of spines below the mandibles. Prothoracic girdle consisting of 8—12
rows of extremely minute spines. Spinal bands well developed at least on the
metathorax and first three abdominal segments, consisting of short pointed spines
with broad bases: these point dorsally or posteriorly.
Two species (oryzae and distorta) show special adaptation to an aquatic environ-
ment: the front spiracles (Fig. 47c) are enormously enlarged and bear at least 75
bulbs: there are patches of bristle-like spines on the locomotory humps (Kriech-
wiilsten) of at least the metathorax and first three abdominal segments.
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 11115)
Group of ambigua Fallen
This group differs from the r/gripes group in the following particulars.
External morphology
The dorsocentrals are stronger and although they decrease markedly in length
forwards, in most species the two pairs of presutural dorsocentrals are still longer
than the acrostichals. Costa reaching only r4 45.
Male genitalia
The basiphallus and mesophallus are generally reduced to narrow strips of chitin
which form a cradle for the large distiphallus. This is similar in form to that of
the nigripes group, but longer than in most species of that group.
Larva
There is a characteristic patch of spines below the mandibles.
Fig. 1. Male genitalia of albipennis Mg. in lateral view, showing the terminology employed.
Fig. 2. Aedeagal apodeme, 9th. sternite etc, of nigripes Mg. in dorsal view, showing the
terminology employed.
AAD — Aedeagal apodeme; HA — Hypandrial apodeme;
AGO — Pregonite; IP — Inner dorsal process of postgonite
AH — Aedeagal hood; MS — Mesaphallus;
BS — Basiphallus; PGO — Postgonite;
DS — Distiphallus; PHO — Phallophore;
| QS — 9th sternite
116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
The species which I can definitely include in the ambigua group are ambigua
Fallen, mobilis Meigen, nigrociliata Hendel, verts Hering, niveipennis Zetterstedt,
ocellaris Hendel, nigrifemur Hendel and yanonis Matsumura. Other species which
probably belong here (but whose larvae and genitalia I have not examined) are
megalopsis Hering and frontosa Becker.
Taken together the nigripes and ambigua group as defined above form a very
natural unit, which contains a large number of species in many parts of the world.
The form of the male genitalia is quite distinct from that of other species of
Agromyza, and this character alone is sufficient to recognise a species as belonging
here. SASAKAWA (1958) has found further characteristic features in the female
genitalia. All the known larvae are feeders on Gramineae.
The only other grass-feeding species of Agromyza are the members of the small
group of cinerascens Macquart, with three known species. These are cinerascens
Macquart, intermittens Becker and /uteitarsis Rondani. This group is not related to
the nigripes and ambigua groups, and indeed in several features stands well apart
from the rest of Agromyza. The genitalia of cinerascens are figured by HERING
(1951b). The phallus is very elongate, quite different from that of most Agromyza
species, though perhaps there is some approach to this condition in Agromyza rubi
Brischke and A. celtidis Nowakowski. The cerci are very large and bear a patch
of spines at their tips. The group is further distinct on various external characters,
as given in HENDEL’s (1931) key. The larvae, however, are still little known, and
their description might throw some new light on the phylogeny of the Agromy-
zinae.
The nigripes group outside the Palaearctic region
The distribution of the nigripes group is probably worldwide. In North America
two species are known (FRICK, 1959). Agr. parvicornis Loew is a pest of maize
(Zea mays L.). The larval mouthparts are figured by PHILLIPS (1914) as having
two large teeth; the hind spiracles of the puparium are only narrowly separated.
This suggests a species closely related to the Palaearctic phragmitidis Hendel,
spenceri sp.nov. and hendeli sp.nov. Frick also records nigripes; this record
however requires confirmation. I suspect from his description and figure that this
species is albipennis.
Many of the species described for Africa by SPENCER (1959 and 1960) belong
here, including penniseti Spencer, susannae Spencer, pallidifrons Spencer,
catherinae Spencer, oliviae Spencer and graminivora Spencer. The life-history of
two of these species, penniseti and graminivora is known. Some of these are close
to Palaearctic species, but they all appear distinct. DE MEIJERE (1940) also
described the larvae of two species from Pennisetum in Java.
COMMENTS ON EARLIER TREATMENTS OF THE GROUP
The following detailed criticism of DE MEIJERE's and HENDEL’s work is neces-
sary in order to clarify the points in which the present treatment disagrees with
theirs, and to explain why I have felt it necessary to modify their conclusions.
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 117
DE MEIJERE in “Die Larven der Agromyzinen” (1925) described larvae of
“nigripes” and phragmitidis. The first was described from Phragmites and refers
to hendeli sp.nov. Both these descriptions are accurate.
In his first supplement (1928) he adds a description of the larva of lucida (as
airae Karl), which is also accurate. The second supplement (1934) includes a
short description of the larva of albipennis. This is erroneous in one important
point: DE MEIJERE interprets the mandibles as consisting of two pairs of alter-
nating teeth: in fact here are four pairs of non-alternating teeth.
In his seventh supplement (1943) he discusses the larvae of the group at some
length and gives a key. This key is vitiated in two respects. First, the mis-
interpretation of the mouthparts of the albipennis larva noted above, secondly a
confusion of the larvae of hendeli (as lucida) and nigripes. This must have
occurred through confusion of the data as these larvae are easily distinguished
from each other; in fact the key contradicts the earlier description of the hendelt
larva (as nigripes in 1925) which was accurate. Also the textfigures accompanying
this article are not properly labelled, and there is considerable doubt as to which
species many of them represent. The result is that there has been a great deal of
confusion over the larval characters of this group, as DE MEIJERE's key has been
the only one hitherto available.
HERING in his “Bestimmungstabellen der Blattminen von Europa (1957) un-
fortunately incorporated DE MEIJERE's key into his work and perpetuated these
errors, as no subsequent studies had been made of the larval morphology of this
group.
The study of this group owes a great deal to HENDEL. However there are two
points on which his treatment in “Die Fliegen der paläarktischen Region” (1931)
requires criticism. First, although the specimens which he used to describe albr-
pennis and nigripes were correctly determined, his use of the relative width in
profile of the cheeks and jowls as diagnostic characters to separate these species
cannot be upheld. The key given is accordingly misleading on this point.
The second point of criticism is more serious. In 1920 he described Agr. lucida
from a swept specimen: later he identified flies bred by HERING from Phragmites
with this species. When writing his later description, HENDEL (1931) describ-
ed one of these specimens, not the type (not unreasonably, as the type is pinned
directly on a long fragile pin and is dangerous to handle). Unfortunately HENDEL
had included two species under the name /zcida, and his 1931 description does not
refer to the same species as the type. Consequently the name has been employed
incorrectly for the Phragmites-feeding species ever since. In this paper the name
lucida is returned to the original species, and the Phragmites-feeding species
redescribed as hendeli sp.nov. I have made HENDEL’s specimens the types of this.
PHYLOGENY
In this assessment of the relationship between the various members of the
nigripes group I have tried to take account of the larval and genitalia characters,
as well as the external morphology of the adult. The best generic characters are in
fact to be found in the larval stages, which are far easier to classify than the adults.
118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
AMBIGUA GROUP
BAETICA N NIGRIPES SPENCERI
DISTORTA ORYZAE GRAMINICOLA PRESPANA ALBIPENNIS LUCIDA PHRAGMITIDIS HENDELI
Graph I. Phylogeny tree of the Agromyza nigripes group
There are two obvious pairs of species. A. oryzae and A. distorta both have
dimorphic puparia, and the larvae are strikingly modified in adaptation to an aquatic
environment. The enormously enlarged front spiracles (Fig. 47c) of these species
are unparalleled elsewhere in the Agromyzidae. The strong dorsocentrals too are
characteristic of both these species. The genitalia and external morphology of the
adult however show that these species are best classified as aberrant members of
the nzgripes group.
Another obvious pair of species is formed by A. graminicola and A. baetica.
Both have identical large antennae (Figs. 23 and 24), and their mesophalli (Figs.
12c and 13c) are modified to form club-shaped appendages. The striking genitalia
modifications of baetica are clearly of recent origin. The separation of these species
must be due to recent geographical isolation.
Three species, phragmitidis, hendeli and spenceri, all produce identical mines on
Phragmites. They have a very characteristic habit of laying a row of eggs on the
leaf, which at once distinguishes their mines from those of other species. The
larval mandibles of all three are identical (Figs. 41a, 42a and 43a), and they all
possess narrow ejaculatory bulbs (Figs. 8a, 9a and 10a), and fairly large disti-
phalli. Therefore in spite of the external differences, I have no doubt that these
three species are closely related.
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 119
The evidence for placing albipennis, nigripes and lucida together is also
strong. A. nigripes and A. lucida are close in respect of their genitalia, and,
although the albipennis genitalia are very characteristic, in external facies and larva
it stands very close to nigripes. All these species have a broad frons (in contrast
to the Phragmites-feeders mentioned above).
A. prespana is not classified so easily. The genitalia of this species (Fig. 11)
are fairly typical of the group, but the larval characters do not allow it to be
closely associated with any of the above groups. The mandibles (Fig. 44a) have
two teeth, but are small, and the hind spiracles (Fig. 44d) are widely spaced and
situated on obvious projections.
We are thus left with four species-groups and A. prespana as species sola.
However the differences which distinguish the oryzae/distorta and graminicola/
baetica groups are obviously much more radical than those separating the rest.
Probably their separation must antedate that of the other groups.
It is also interesting to note which characters prove to be of no phylogenetic
value. The colour of the squamal fringe is a good specific character, but of no
generic importance. The antennae too are enlarged in several not closely related
species (viz., hendeli and albipennis/nigripes): clearly a tendency to variation in
the shape of the antennae is part of the ancestral genetic make-up of this group.
In classifying the larvae too we must take account of the shape of the mandibles,
not merely the number of teeth. Six species have larvae with two teeth, but there
are differences in the shape of the mandibles. The three Phragmites species,
phragmitidis, hendeli and spenceri, have large broad mandibles: those of grami-
nicola and baetica are narrower, and those of prespana smaller still. Probably two
was the original number of teeth for the group, and its retention therefore not
necessarily indicative of close affinity. The large number of teeth shown by two
species-groups would then represent an innovation. The fact that DE MEIJERE's
Javan species (1940) and the American A. parvicornis Loew (PHILLIPS, 1914)
also possess two teeth lends support to this view.
HOST RELATIONSHIP
A balanced assessment of the degree of host specialisation in this group is
hampered by two difficulties. First the botanists’ ideas on the classification of the
Gramineae are not yet very stable. Secondly our knowledge of the host-range of
Agromyzids is largely biased in favour of the more easily recognised grasses, such
as Phragmites, Glyceria, Phalaris and Deschampsia. Accurate records from other
hosts are relatively meagre, but this is clearly more the fault of the entomologists
than the flies.
However, regarding these four hosts, one can speak with reasonable certainty.
The first striking fact is that Phragmites is the host of five monophagous species.
This is clearly good circumstantial evidence for botanical systems which make the
Arundineae a separate tribe. Of the other three hosts, Glycerza and Deschampsia
share two species, neither of which occurs on Phalaris, while albipennis, which is
abundant on Phalaris and occurs on many other grasses (particularly the Hordeae),
never occurs on Glyceria, though it was once bred from Deschampsza. Probably
120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
we should be unwise to base any argument on the host-selection of albipennis
which is clearly very catholic in its tastes: but the fact that /zcida is common on
these two plants and rarely occurs on any others does seem of significance. We
thus have a priori grounds for welcoming a possible connection between the Gly-
cerieae and Aveneae, but they are admittedly not based on very strong evidence
in the present state of our knowledge of this group.
The following list uses the classification given in CLAPHAM, TUTIN and WAR-
BURG (1952).
* recorded from this host once only.
Oryzeae | Oryza sativa L. oryzae (Munakata)
| o a 6 o 5
| Zizania latifolia Turcz. oryzae (Munakata)
Arundineae | Phragmites communis Trin. baetica sp.nov.
graminicola Hendel
hendeli sp.nov.
phragmitidis Hendel
Spencer? sp.nov.
Glycerieae | Glyceria maxima (Hartm.) distorta Griffiths
| Holmb. lucida Hendel
| nigripes Meigen
| Glyceria fluitans (L.) R.Br. nigripes Meigen
Festuceae | Festuca sp. *albipennis Meigen
| Poa spp. albipennis Meigen
| Echinaria capitata Desf. *lucida Hendel
Brachypodieae Brachypodium sylvaticum nigripes ssp. brachypodii
| (Huds.) Beauv. ssp. nov.
Hordeae Triticum aestivum L. prespana Spencer
| *albipennis Meigen
Secale cereale L. *albipennis Meigen
| Hordeum spp. albipennis Meigen
Aveneae | Arrhenatherum elatius (L.) | *albipennis Meigen
J, 6 CG, Jel
Holcus spp. nigripes Meigen
| Deschampsia caesp'tosa (L.) lucida Hendel
Beauv. | *albipennis Meigen
*nigripes Meigen
Agrosteae | Agrostis stolonifera L. *albipennis Meigen
Agrostis canina L. *nigripes Meigen
Phleum pratense L. | *albipennis Meigen
Phalarideae Phalaris arundinacea L. | albipennis Meigen
Phalaris erucaeformis L. albipennis Meigen
Paniceae Setaria viridis Beauv. *albipennis Meigen
I note that in many classifications the Seslerieae (including Echinaria) are
erected as a separate tribe and placed next to the Aveneae. This certainly makes
the record of Jucida from that plant look more reasonable: [cda obviously has a
narrow range of hosts.
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 121
There are many references in the literature on Applied Entomology to “Agro-
myza nigripes” occurring as a pest of cereal crops. Unfortunately very few of the
specimens concerned have been examined by specialists, and I have thought it best
to disregard these records for the purpose of this paper. The available evidence
suggests that these records will mainly refer to A. albipennis, which is clearly the
commonest species on the Hordeae. This species has been confirmed on barley in
Japan and Germany, and on wheat in Great Britain. The other species which has
occurred on wheat is the recently discovered A. prespana, which may possibly be
found in Britain. References to “Agr. nigripes’ having only one generation a
year might well apply to this species. The record of true A. nigripes has not been
confirmed from cereal grasses.
Discussion of the host selection of the individual species in cases where oligo-
phagy occurs will be found under the species concerned.
KEYS FOR IDENTIFICATION
ADULTS!)
1. 5 of 6 pairs of dorsocentrals clearly developed as bristles, exceeding the height
of the acrostichals in profile (1 or 2 pairs being presutural). a with short
white pubescence. On aquatic grasses . . IRE SOR 2
— At most only the four postsutural omeen ded ee as bristles in
proniles byes bate on weakly pubescent. nnen a ARCS
2. Frons broad, 11/,—2 times the eye-width; 2—3 strong lower orbital bristles
(Fig. 23). Ocellar bristles reaching as far as the base of the 2nd upper
orbitals. Acrostichals 8-rowed. 4: ejaculatory bulb (Fig. 14a) small, transpa-
rent in distal 1/,, the central channel reaching 2/3 of its length; thorn-like
process of the left side of the mesophallus (Fig. 14c) very strongly developed.
Europe. On Glyceria maxima. . . . + distorta Griffiths
— Frons narrower: normally 3—5 short louer Sel redes (Fig. 26). Ocellar
bristles very short, not reaching as far as the base of the 2nd upper orbitals.
Acrostichals 6-rowed. 4: ejaculatory bulb (Fig. 15a) larger, moderately
sclerotised except along its distal edge; central channel reaching nearly the
distal edge; mesophallus without thorn-like process. Siberia and Japan. On
Oro zaand 0101 NEN ENEN oryzae (Munakata)
3. Squamal fringe white . . . Ph A Pa
—- Squamal fringe normally adhere hars nb ne D ch agent Normally
only 4 orbital bristles (Fig. 17). Frons broad, 1.2—1.8 times the eye-width.
3rd antennal segment somewhat broadened apically: nigripes Meigen?) . 8
== Famomik ia (Ade PE Mene been Goce Wiebe! CA O
4. 3rd antennal segment small and rounded (Fig. 21); 5 orbital bristles, usually
1) Excluding /unulata Sasakawa.
2) A very common, variable species. If squamal fringe darker (as often in Scandinavian
specimens) it may be confused with /ucida or hendeli; the former is distinguished by the
shape of the antennae: the latter by the narrower frons and more numerous orbital bristles.
hake
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
fairly weak. Narrow frons (0.8—1.2 times the eye-width). 4: ejaculatory
bulb (Fig. 10a) dark and narrow. On Phragmites. . . phragmitidis Hendel
3rd antennal segment larger. G': ejaculatory bulb broader and paler. . 5
. 3rd antennal segment very much enlarged (Figs. 23 and 24). At least the
first pair of lower orbital bristles directed more or less backwards. &:
ejaculatory bulb not so broad: mesophallus (Figs. 12c and 13c) developed
into a pair of club-shaped appendages. . . NE €
3rd antennal segment not enlarged so much (Ga. 16). ora orbitals all
directed more or less inwards. g': ejaculatory bulb (Figs. 3a and 4a) broad;
mesophallus not as above: albipennis Meigen. . . . . a e
. 4—5 orbital bristles (Fig. 23). Frons narrow, 1—1.3 times the eye-width.
Eyes bare. g': 9th sternite symmetrical: aedeagal apodeme normal. Northern,
central and eastern Europe. On Phragmites. . . . . graminicola Hendel
5—6 orbital bristles (Fig. 24). Frons broader, 1.5 times the eye-width. Eyes
with short pubescence. Cheeks and orbits clearly visible in profile. g'; 9th
sternite asymmetrical (Fig. 13d), the left side about twice as long as the
right; aedeagal a ci broadened posteriorly. Southern Spain.
On Phragmites . . . . baetica sp.nov.
. d': ejaculatory bulb (Fig. sa) fa les ee Common species. On
various grasses, especially Phalaris . . . . . . albipennis Meigen s.s.
d': ejaculatory bulb (Fig. 4a) very short, much wider than high, with a
a black central channel which bifurcates at its tip. Finland.
albipennis fennica subsp. nov.
Wing lot 2.0—2.8 mm. d': eee bulb (Fig. 5a) with its distal edge
at least partially black. Common species. On Glyceria, Holcus and other
PASSES Beer nr NESS
Wing-length 1.8—2.2 mm. &: oe bulb (Fig. 6a) mostly transparent,
but with an obvious black patch at its base on one side. Corsica. On Brachy-
DOU 6 ONE . + + . migripes brachypodi subsp. nov.
. 3rd antennal nn edie dimorphic (Figs. 22 and 22a), in the female
longer than high with a short fringe of upturned hairs, in the male larger,
with a very long fringe. Antennae pointing downwards, the frons projecting
above them. Facial keel clearly projecting in profile. On Triticum .
prespana Spences
- 3rd antennal segment without such a fringe. Facial keel not ES ne An-
tennae not markedly pointing downwards. . . . 10
. 3rd antennal segment enlarged (Fig. 19). 5—6 strong ol nes directed
more or less backwards. Frons narrow, 0.8—1.2 times the eye-width. &':
ejuculatory bulb (Fig. 8a) narrow and black. On Phragmites .
hendeli sp.nov. (= lucida Hendel 1931 p.p. nec 1920)
3rd ential segment small and rounded, 4—5 orbital bristles. . . . 11
4th costal segment short (Fig. 31): ratio of 2nd to 4th costal segments 4—4.2.
Face-line very flattened (Fig. 20). 8: characteristic enlarged distiphallus
(Fig. 9b). Alpine species. On Phragmites. . . . . . spenceri sp.nov.
|
|
cn
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 123
4th costal segment longer (Fig. 29): ratio of 2nd to 4th costal segments
2.7—4.0. Face-line not so obviously flattened (Fig. 18). g': distiphallus
(Fig. 7b) of moderate size, rather elongate. Widespread. Mainly on Des-
champsia and Glyceria . . . . . . . lucida Hendel (= airae Karl)
LARVAE
. Mandibles (Fig. 47a) with 5 (rarely 4) alternating teeth. Front spiracles
(Fig. 47c) large and circular with at least 75 bulbs. At least the metathorax
and first three abdominal segments with thick patches of bristle-like spines
ventrally on the locomotory humps (Kriechwiilsten). On aquatic grasses . 2
Mandibles with 2—4 teeth, not or only slightly alternating. Front spiracles
small, with less than 25 bulbs. No special spines ventrally. . . . . 3
. Front spiracles with 75—160 bulbs. Patches of bristle-like spines on the
locomotory humps present on mesothorax, metathorax and all abdominal
segments. Japan and Siberia. On Oryza and Zizania. . oryzae (Munakata)
Front spiracles (Fig. 47c) with 200—250 bulbs. Patches of bristle-like spines
on the locomotory humps present only on metathorax and first three abdominal
segments. Europe. On Glyceria maxima . . . . . . distorta Griffiths
. Mandibles with 2 teeth . . 4
Mandibles (Fig. 40a) with 3 ch, esami halle ‘afbeet 6th to “sth
abdominal segments with clear bands of spines laterally. Mainly on Glyceria
and Deschampsia . . cda plendel (Saer Karl)
Mandibles with 4 teeth (Eis 38a and 39a). Mesothoracic “ea represented
by at least a patch of spines ventrally . . . . . io DCS
. Hind spiracles separated by more than twice their own Felis Be laid singly
mearsthesedoezotitherlear 2 20: 5
Hind spiracles separated by less than twice ene own à la Bee md smelly
in a row of 3 or 4, the resulting larvae producing a communal mine.
Oy PATES EX REMO OI ven mnie IA EE
. Hind spiracles only on small projections. On Phragmites. . . 6
Hind spiracles (Fig. 44d) on large Keen Mandibles (ie 44a) re-
latively small. On Triticum. . . |. . prespana Spencer
. Adjacent bulbs of the hind spiracles (Fig 46d) forming an angle of 80°.
Pupation outside the mine. Southern Spain. . . . . baetica sp.nov.
Adjacent bulbs of the hind-spiracles (Fig. 45d) ut an angle of 90°—
100°. in the mine. Northern, central and eastern Europe .
graminicola Hendel
. Processes si the aise a (Fig 42b) consisting of transparent
CHINE . + phragmitidis Hendel
Processes of Be arde all ragno (LE 41b and ee ae of
mainly brown and black chitin .
hendeli sp.nov. (= (ede He dl 1931 DI nec 1920)
and spenceri sp. nov. (Alpine species).
(No larvel preparation of spenceri was available for close comparison).
124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
8. Hind spiracles (fig. 39d) contiguous: their adjacent bulbs forming an angle
of 80°. On various grasses, especially Phalaris . . . albipennis Meigen
— Hind spiracles (Fig. 38d) separated by 11/,—2 times their own width: their
adjacent bulbs forming an oe of 100°. On Glycerza, Holcus and other
OSES 5 : . . . . nigripes Meigen
A. nigripes DD Ee nov. was Dal da err in Corsica. Larval
morphology not known.
DESCRIPTIONS
Agromyza albipennis Meigen 1830
Agromyza albipennis Meigen, 1930, Syst. Beschr. bekann. eur. zweifl. Insekt,, vol. 6, p.
lye
Agromyza albo-hyalinata Zetterstedt, 1848, Dipt. Scand., vol. 7, p. 2742. Syn. nov.
Agromyza nigripes Hendel (nec Meigen) pro parte, 1920, Arch. Naturgesch., vol. 84,
9, 31112,
lexthieures M sans besc 116527, 393,396, 3975394948)
The Zetterstedt species albo-hyalinata is synonymised as a result of my examin-
ation of the g genitalia of the type.
Frons of the broader type, 1.2—2 times the width of an eye at the level of the
front ocellus; not or only slightly narrowing forwards. Antennae a little separated
at their bases. In profile (Fig. 16) the cheeks and orbits usually clearly visible as
a narrow ring around the eye, often slightly projecting at the level of the antennae.
Ratio of the eye. height to the height of the jowls usually 3.5—5 (though
occasionally the jowls may be narrower, the ratio being as much as 11). 4—5
orbital bristles (one specimen has 6 on one side only). Upper and lower orbitals
clearly distinguished, the later directed inwards or inwards and backwards. When
a third lower orbital bristle is present this is usually small. Third antennal segment
somewhat broadened apically with a rounded tip. Colour of head variable: usually
at least the first antennal segment pale; sometimes the frons and the first two
antennal segments yellowish brown.
6—8 rows of acrostichals. Legs variable in colour. Usually at least the front
femora yellow at their tips. Often the tips of all femora, and the front tarsi
yellowish. Sometimes however the legs are completely black.
Ratio of 2nd and 3rd costal segments (Fig. 27) 2.5—4.0; 2nd and 4th 3.5—5.
Wing tip usually between the ends of ry 45 and m, ‚>. All wing veins, except
sometimes the costa, yellowish, especially towards the base of the wing. Membrane
transparent or slightly whitish tinged. Squama white with a white fringe, only its
border being sometimes yellowish. Wing length 2.0—3.0 mm.
3 genitalia. Distance from the end of the aedeagal apodeme to the end of the
aedeagal hood about 1 mm. The aedeagal apodeme is conspicuously thickened at
its anterior end and slightly bent downwards over its length. The strengthened
part posterior to the attachment of the inner processes of the postgonites occupies
about 1/; of the whole apodeme. The 9th sternite is long, strongly bowed down-
wards especially towards the tip. Hypandrial apodeme not clearly differentiated,
being more ot less continuous with the 9th sternite. The tip of the hypandrial
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 125
apodeme reaches 2/3 of the length of the aedeagal apodeme. Pregonites as shown
for nigripes (Fig. 2) though relatively smaller. Postgonites small, more or less
club-shaped, similar to those of nigripes (Fig. 2). Aedeagal hood weakly chitinised;
its width about twice that of the posterior end of the aedeagal apodeme. Phallo-
phore as figured (Fig. 1).
Basiphallus (Fig. 3c) very long and moderately slender, consisting of two rods,
both clearly defined, the right rod somewhat stouter than the left. Mesophallus
consisting of two small triangular plates which are narrowly separated at their tips
where they meet beneath the distiphallus. The distiphallus (Fig. 3b) is more
weakly bent than in any other species, its rear part not bent downwards. Disti-
phallus clearly shorter than the basiphallus.
The ejaculatory bulb (Fig. 3a) is large and scallop-shaped, clearly broader than
long. Sometimes the medial channel is visible. Its sclerotisation is moderately
strong. Stem short and broad.
Larva. Antennae very small; maxillary palpus large and thick: slender but
strong longitudinal sclerite. Narrow mandibular abductor apodeme present.
Mandibles (Fig. 39a) rather narrow, with 4 teeth, the lower two smaller; usually
an obvious protuberance posteriorly behind the point of attachment of the labial
sclerite. 2—3 pairs of inconspicous filaments above the mandibles.
Labial sclerite (Fig. 39b) and paraclypeal phragma slender, but strongly
sclerotised. Small antero-dorsal process at base of paraclypeal phragma.
Head without spines. Prothoracic girdle consisting of about 8 rows of minute
spines. Mesothoracic girdle present only ventrally, with about 4 irregular rows of
spines. Metathorax to 4th abdominal segment with broad bands of large spines
pointing postero-dorsally; metathorax with 6—7 rows of spines, lst to 3rd ab-
dominal segments 8—10 rows, 4th abdominal segment 6 rows; all bands weaker
dorsally. 5th abdominal segment with a well developed band at least laterally,
with 3—5 rows of spines. Following segments bare laterally, with a few spines
ventrally in 2 rows. Also numerous small sharp spines in the anal region.
Front spiracles (Fig. 39c) with 12—16 bulbs, consisting of two almost equal
horns on a long somewhat twisting atrium. Hind spiracles (Fig. 39d) contiguous:
their bulbs oval-shaped, of moderate size; adjacent bulbs forming an angle of 80°.
Length (full-grown) 3.75—4.5 mm.
Biology. Mine (Fig. 48) on upperside of leaf. Eggs laid singly near the
edge of the leaf, usually some distance from the leaf tip. The larva at first makes
a narrow linear mine towards the leaf tip, then turns and produces a broad
irregular channel, eating towards the base of the leaf. Frass irregularly spaced, in
large particles. In fresh mines the zig-zag pattern of the feeding lines is often
visible.
Pupation follows either inside or outside the mine: if outside, the puparia adhere
to the leaf near the mine. At the Ash Vale locality (Surrey) about 50% of the
larvae were observed to pupate in the mine. The colour of the puparia varies from
black to red.
There are two or probably often three generations. Adults are found first in
126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
June and continue to early September. Mines from June to October, but especially
common in early September.
Material examined. In the author's collection. &, 2 2 Woodwalton Fen, Hunts.
17—22.VIII.60, ex Phalaris arundinacea: emerged 29.VIII, 3.IX, 30.11.61. 2 Woodwalton
Fen, Hunts., 22.VIII.60 ex Arrhenatherum elatius: emerged 29.VIII. 3 4, 9 2, swept from
Woodwalton Fen, Hunts., 16—25.VIII.60 (4 preparations made with Woodwalton material).
3 ê, 3 2, Ash Vale, Surrey, 5.IX.53, ex Phalaris arundinacea: emerged Sept. 53 and
May-July 54. 4, 6 ®, Ash Vale, Surrey, 15.VIII.54 ex Phalaris arundinacea: emerged
August/Sept. 54 and June 55, ® swept at Ash Vale, Surrey, 5.IX.53. 10 &, 16 9, Ash
Vale, Surrey, 5.IX.60, ex Phalaris arundinacea: emerged March/April 61 (4 preparations
made from Ash Vale material). 4 (with preparation), ® Rickmansworth, Herts., ex Pha-
laris arundinacea 17.1X.53: emerged 13.V.54. 9 Finchley, Middx., 31.VIII.53, ex Phalaris
arundinacea: emerged 17.XI.
Larval preparations from Ash Vale, Surrey, 5.IX.60 (5 preparations) and Woodwalton
Fen, Hunts., 16.VIII.61 and 29.VIII.61 (3 preparations) (all from Phalaris arundinacea).
In K. A. SPENCER’s collection. & (with preparation), Oxford, 2.X.52: emerged 5.V.53
ex Phalaris arundinacea.
In the HENDEL collection (Vienna). 9 ex Hordeum muralis, Wien. © Prater (Wien).
6 (with preparation) labelled ”T” in red (No data: presumably Vienna. Probably the
specimen used by Hendel for his description in Hendel (1931)). ® Hortobágy leg.
Kertész. 9, Madara, 928, leg. Santády, VI.30, 4, Silesia, leg. Kertész. 9 Südtirol, VIII. 4
Ebenew, 1.VIII.81, leg. Becker. 3 4, 2 9, no data (presumably Vienna area). & (with
preparation), Montserrat, Spain, col. Strobl. & (with preparation), Böhmerwald, Rachel,
24.VIII.15, leg. Zerny.
In the GROSCHKE collection (Stuttgart). 9, 3.VII.49, ex Setaria viridis, Mecklenburg,
Schwaan, Buhr, 63.
In the HERING collection (Berlin), 2 ®, ex Secale cereale, Mecklenburg, Gr. Lüsewitz
168 & 169, 16.VII.51, leg. Buhr. 9 ex Hordeum vulgare, 16.VII.49, Mecklenburg, Rostock,
Buhr, 499. 4 9, ex Phalaris arundinacea Grünhof b. Stettin, leg. v. Oethingen. 9 Mecklen-
burg, Rostock, Buhr, 16.VII.49 (swept). 9, ex Agrostis alba,, 16.VII.49, Mecklenburg,
Rostock, Buhr, 435. ®, ex Deschampsia caespitosa, 16.VII.49, Mecklenburg, Rostock, Buhr,
462. 9, ex Phleum pratense, 9.VII.49, Mecklenburg, Rostock, Buhr, 483. & (with
preparation), ex Poa trivialis, 8.VII.49, Mecklenburg, Rostock, Buhr, 487. 4, ex Poa sp.
13.VII.49, Mecklenburg, Schwaan, Buhr, 275. ®, ex Festuca sp., 12.VIL49, Mecklenburg,
Rostock, Buhr, 402. & (with preparation), ex Phragmites communis, Verson b. Caen,
21.VII.42, Buhr, 79. &, swept at Berlin, Finkankg. 6.V.29. Also a larval preparation from
Lüsewitz, 30.III.32, No. 5884, ex Phalaris arundinacea.
In the Rypen collection (Hälsinborg, Sweden). 9 Ang. 23.VII.33. ® Vg. Torpa
22.VI.46. 4 (with preparation), Gotland, Etelheim, 19.VII.50. 4 Gotland, 15.VII.50. 4
(with preparation) Gotland, Bogeviken Slite, 15.VIII.50. 4 4, 2 ®, Närke Mulhytten tf.
Lillan, 4—5.VII.52. 2 6, 4 9, Skane, Skälderviken, 5.VIII.55. 9 Skane, Sköralid, 18.V.48.
9 Skane, Bölinge, 16.VI.55. Q, Kullaborg, 20.VIII.40. 9, Angelh., 15.VI.50. 9, Väster
Gotland, Ulricehamn, 14.VII.47. @ Ängermanland, Docksta, 18.VI.54.
In the ZETTERSTEDT collection (Lund). ¢ (with preparation), type of the synonymous
albo-hyalinata Zett. from Esperöd. The differences noted by Zetterstedt in his description
are due to the specimen having been killed soon after emergence before its colours had set.
There are also accurate records for Holland (DE MEIJERE 1934) and Scotland
(SPENCER 1956). SASAKAWA (in litt.) knows this species from Japan.
Distribution. Great Britain (incl. Scotland), France, Northern Spain,
Holland, Sweden, Germany, Austria, North Italy, Hungary, Czechoslovakia, Po-
land, Japan. Two © from Kamtschatka were referred here by HENDEL (1931). I
cannot be certain if these are albzpennis. The matter must rest until a & is
obtained.
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 127
Figs. 3—4. Male genitalia of Agromyza: 3, albipennis Meigen sensu stricto, 4, albipennis
fennica subsp. nov.: a — ejaculatory bulb; b — lateral view of phallus or distiphallus;
c — ventral view of phallus
The species is generally common. After the ubiquitous nigripes it is the next
commonest species to turn up in swept material.
Host plants. Gramineae, tribes Festuceae, Hordeae, Aveneae, Agrosteae,
Phalarideae and Paniceae.
The species is always abundant on Phalaris arundinacea L. Many of the records
from other genera have only been made on one occasion, and many of them may
therefore only be occasional foodplants (particularly the record from Deschampsia
from which /ucida has been bred many times but albipennis and nigripes once
only). I therefore now give a separate list of the known normal food-plants of
this species from those from which it has been bred on one occasion only.
Normal food-plants: Festuceae, Poa (bred from P. trivialis L. and Poa sp.).
Hordeae, Hordeum (bred from H. vulgare L. and H. murinum L.). Phalarideae,
Phalaris arundinacea L. (very common here).
Recorded as food-plants once only: Festuceae, Festuca sp. Hordeae, Secale
cereale L. Aveneae, Arrhenatherum elatlius (L.) J. & C. Presl. Deschampsia
caespitosa (L.) Beauv. Agrosteae, Agrostis stolonifera L. (= alba) Phleum pra-
tense L. Paniceae, Setaria viridis Beauv.
There are several references in the British literature (by SPENCER and myself)
to this species being bred from Phragmites. These are all due to confusion of the
food-plant, which was Phalaris, not Phragmites. There is one male in HERING’s
128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
collection (listed above) labelled Phragmites collected by Dr. BuHR. He cannot
however trace the mine from which it was bred, so that I regard this record as
doubtful. In any case Phragmites cannot be a normal food-plant: observation in
this country has shown that where Phragmites and Phalaris grow together, albi-
pennis occurs in profusion on Phalaris, but is never found on Phragmites.
Records of this species on G/yceria are inaccurate and refer to nigripes.
Agromyza albipennis Meigen fennica subspec. nov.
Textfigure 4a.
Frons broad, 1.8 times the width of an eye at the level of the front ocellus.
Cheeks and jowls clearly visible in profile. Eye small, ratio of eye-height to the
height of the jowls 3.0. 3 strong lower orbital bristles.
Wing-veins brown; membrane whitish tinged with fairly obvious pubescence.
Squamal fringe dull white. Wing-length 1.8 mm.
Not distinguishable on external characters from small examples of the typical
form.
3 genitalia. Aedeagal apodeme, 9th sternite, postgonites, pregonites and aedea-
gal hood as for the typical form. Basiphallus consisting of two long rods with an
obvious ventral appendage composed of transparent chitin (in typical albipennis
this is vestigial). Distiphallus as in the typical form: distiphallus and basiphallus
of equal length.
The ejaculatory bulb (Fig. 4a) much smaller than in the typical form; much
broader than long with a strongly sclerotised central channel which bifurcates at
its tip. The stem is very large (contrast the typical form).
Larval stage and biology unknown.
Material examined. In the HENDEL collection. Type g (with prepa-
ration), Messuby, coll. FREY.
I associate with this specimen 2 9 labelled Kuusamo and Kuopio, coll. FREY.
Distribution. Finland.
The genitalia of this subspecies are strikingly distinct from the typical form of
albipennis. However the external morphology is not incompatible with that of
small specimens of albipennis, and the distiphallus is of the characteristic albi-
pennis type. I have accordingly given the form only subspecific rank.
Agromyza nigripes Meigen 1830 pro parte
Agromyza nigripes Meigen, 1830, pro parte, Syst. Beschr. bekann. eur. zweifl. Insekt.,
vol. 6, p. 170. (mec Zetterstedt, Schiner, Rondani, Stary pro parte et passim).
Agromyza nigra Zetterstedt, 1840, Ins. Lap., p. 788.
Agromyza carbonaria Zetterstedt, 1848, pro parte. Dipt. Scand., vol. 7, p. 2739, et 1860,
loc. cit., vol. 14, p. 6451 (var. b duplo fere minor): etiam Schiner, Bezzi et passim.
Agromyza agrosticola Hering, 1927, Z. angew. Ent., vol. 13, p. 191.
Textfigures 2, 5a, 5b, 5c, 17, 28, 38a, 38b, 38c, 38d, 38e, 49
A fuller account of the synonymy is given by HENDEL (1931). There is one
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 129
small deletion to make. The specimen of Agr. holosericea Strobl. nec Bouché is
lucida Hendel.
Frons of the broader type, 1.2—1.8 times the width of an eye at the level of
the front ocellus, parallel-sided. Antennae clearly separated at their bases. In
profile (Fig. 17) the cheeks and orbits are usually visible at least narrowly. Ratio
of the eye-height to the height of the jowls 3.5—7, though occasionally the jowls
are narrower, the ratio being as much as 11. 4 strong orbital bristles (rarely a
small 5th), the lower pair usually directed inwards and backwards. 3rd antennal
segment slightly broadened apically, rounded at its tip. Colour of head usually
completely black; occasionally the frons and first two antennal segments dark
brown.
Acrostichals in 6— 8 rows. Legs completely black, or the tips of the front femora
only yellow.
Ratio of 2nd and 3rd costal segments (Fig. 28) 2.7—3.4; 2nd and 4th 3.3—5.
Wing-tip between the ends of ry 45 and m, ‚5, rarely nearer the former. Wing
veins brown, membrane obviously brownish tinged. Squamal fringe usually
ochreous or brown, but varying from almost black to greyish white (in particular
most Scandinavian specimens have a dark-brown to blackish fringe). Wing-length
2.0 —2,8 mm.
d' genitalia. Distance from the end of the aedeagal apodeme to the end of the
aedeagal hood 0.8 mm. Aedeagal apodeme thick, heavily sclerotised, the strength-
ened part posterior to the attachment of the inner processes of the postgonites
occupying 1/5 of the whole apodeme. 9th sternite short and rounded, bearing a
narrow hypandrial apodeme which occupies 1/, of the length of the whole
sternite, its tip reaches a little over 1/, of the length of the aedeagal apodeme.
Pregonites large, postgonites small (Fig. 2). Aedeagal hood moderate-sized,
strongly sclerotised around its edges. Phallophore as shown for albipennis (Fig. 1).
Basiphallus (Fig. 5c) consisting of two strongly sclerotised rods, the right rod
broader than the left. There are small ventral appendages composed of transparent
Fig. 5. Male genitalia of Agromyza nigripes Meigen s.s.: a — ejaculatory bulb; b — lateral
view of distiphallus; c — ventral view of phallus; d — an aberrant ejaculatory bulb shown
by one Viennese specimen
130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
7C
Figs. 6—7. Male genitalia of Agromyza: 6, nigripes brachypodii subsp. nov.; 7, lucida
Hendel: a — ejaculatory bulb; b — lateral view of distiphallus; c — ventral view of phallus
view of phallus
sclerotin. Mesophallus consisting of two plates, coming to a point where they al-
most meet beneath the distiphallus. Distiphallus (Fig. 5b) with its rear part
strongly bent downwards; usually also slightly bent at its tip (compare hendel).
Size of distiphallus variable: usually at least slightly longer than the basiphallus,
sometimes very much so.
Ejaculatory bulb (Fig. 5a) small and weakly sclerotised; central channel usually
clearly visible. Its distal edge characteristically black marked, at least centrally, in
most examples. Stem short.
This species shows considerable variation in the colour of the squamal fringe.
Generally it is darker in the males, especially from Scandinavia. In large females
it may be almost white. Also the phallus is rather variable; many specimens from
Central Europe have a very large distiphallus and broader rods to the basiphallus.
However intermediates are common, and variation is often very marked even in
specimens from the same locality.
One Viennese specimen has a very unusual ejaculatory bulb, but is otherwise
normal. This is shown in Fig. 5d. I regard this specimen merely as an aberration
of nigripes.
Larva. Antennae small; maxillary palpus large and thick; slender longitudinal
sclerite only weakly sclerotised. No mandibular abductor apodeme. Mandibles
(Fig. 38a) rather narrow, with 4 teeth, the lower two smaller; a protuberance
posteriorly beneath the point of attachment of the labial sclerite. 3 pairs of
filaments above the mandibles.
Labial sclerite (Fig. 38b) slender and straight. Paraclypeal phragma slender but
strongly sclerotised; its dorsal and median processes fused apically; small antero-
dorsal process at its base.
Head without spines. Prothoracic girdle consisting of about 8 rows of extremely
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 131
minute spines. Mesothoracic girdle very reduced, consisting only of a patch of
small irregular spines ventrally. Methathorax and first five abdominal segments
with continuous bands of irregular dorsally and posterodorsally directed spines
(Fig. 38c) in 7—9 rows; all spines smaller than in alb/pennts. 6th abdominal seg-
ment with a weaker, irregular band of spines in about 5 rows laterally and ventral-
ly; bare dorsally. 7th and 8th segments with about 4 rows of slender pointed spines,
mainly ventrally. Also anal region with numerous small pointed spines.
Front spiracles (Fig. 38c) small, with two horns, the anterior being clearly
longer than the posterior; 16—20 small bulbs in two rows; atrium long, with
several kinks. Hind spiracles (Fig. 38d) separated by 115—2 times their width;
their bulbs oval-shaped, rather widely separated from each other; adjacent bulbs
forming an angle of 100°.
Length (full-grown) 3.75—4.5 mm.
Biology. Mine (Fig. 49) on upperside of leaf. Eggs laid singly, usually on
the edge of the leaf or near the tip. The narrow early channel of the larva usually
swallowed up by its later broader mine. Frass particles fairly conspicuous, irregu-
larly scattered. The mine is deep; on G/yceria parts of the lower parenchyma are
consumed, but the larva generally does not cross the midrib, forming its mine on
one side of the leaf.
Pupation follows outside the mine, the puparia adhering to the leaf near the
mine. Puparia usually black, sometimes reddish.
Two generations. Flies in May/June and again in August/early September.
First generation of larvae in June, second in late August to September.
Material examined. In the author’s collection. & (with preparation) Ash Vale,
Surrey, 5.IX.53, ex Glyceria maxima: emerged 15.VI.54. 6 & (with 2 preparations), 7 9,
Ash Vale, Surrey, 5.IX.60, ex Glyceria maxima: emerged Sept. 60 and March/April 61. 9,
Beaconsfield, Bucks. 6.1X.53, ex Holcus lanatus: emerged 29.1V.54. 6 2, 9, swept at
Woodwalton Fen, Hunts. Aug./Sept. 60. 2 2, swept from Chippenham Fen, Cambs.,
5.VI.55. @ swept at Bookham, Surrey, 23.V.54.
Larval preparations from Ash Vale, Surrey, 5.IX.60, ex Glyceria maxima (two prepara-
tions) and Woodwalton Fen, Hunts., 25.VIII.60, same host.
In K. A. SPENCER’s collection. 9 Chippenham Fen, Cambs., 4.VII.58, ex Deschampsia
caespitosa: emerged 11.VII.58.
In the HENDEL collection. 2 & (with preparations), ®, Messuby, Frey. & (with
preparation), 2 9, Pojo, Frey. 2 4 (with preparations), 2 2 Karislojo, Frey. 2, Kuopio,
Frey. 2, Summatti, Frey. 2 & (with 1 preparation), Hammern, Ost. sup., leg. Mik, 6.VIII.83.
©, Hammern, Ost. sup., 10.IX.72, leg. Mik. 2, Linz, Ost. inf., 4.VII.67, leg. Mik. & (with
preparation), Hainfeld, Ost. inf., 21.VII.97, leg. Mik. 2 & (with preparations), Freistadt,
Ost sup., 26.VI.68, leg. Mik. 4 (with preparation), Baarn, Holland, 14.VI, leg. de Meijere.
4 Felsöhägi, leg. Méhely. 2, Körösmezö, leg. Kertész, July 1917. & (with preparation),
Tisawitz (?), Aug. 1919. & (with preparation), Caltenlichtgeben, Ost. inf. August. 4 4
(with preparations), @ Ossiach, Kärnten, July/August, leg. Hendel. 2 ¢ (with preparations),
Hoheneich, Ost. inf. leg. Zerny. 4 (with preparation), Almsee, Ost. sup., leg. Czerny. 4
(with preparation), 2 9, Prerbaum, Ost. inf, 1.VI.17. 4 (with preparation), 2 Q Langenzers-
dorf, Ost. inf. July. 2 & (with preparation), Prater, Wien, May col. Hendel. 6 & (with 4
preparations), 6 ® without data, presumably Vienna region, leg. Hendel (including a speci-
men labelled ”T” in red, presumably the specimen used for the description in Hendel (1931) ).
In the GROSCHKE collection. 2 9, Baienfurt, Württemburg, 19.VI.55, leg. Groschke.
4 2, Stempeda, Harz, 19 & 23.VI.27, Nos. 3161 & 3142, Hering.
132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
In the HERING collection, 9, Güntersberg an Oder, 5.IX.24, ex Glyceria maxima, Hering
2600. & (with preparation), Maribo, Denmark, 4.V.32, ex Glyceria maxima, leg. Sönderup.
Q, Prerow (Darss.), July 1926, ex Agrostis canina: emerged 24.VIII, Hering, 2978 (type of
the synonymous agrosticola Hg.). 4 &, 2, Stempeda, Harz, 19—27.V1.27, Hering, nos. 3141,
3142, 3148, 3161. 2 & (with 1 preparation), Krippen, Sachs. Schweiz, 27.V1.37, leg. Hering.
In the RYDéN collection. 4 & (with 1 preparation), 4 9, Vg. Ulricehamn, June/July 46—
48. & (with preparation), 9, Bohuslän, Ljungskile, 9 and 11.VIII.51. 9, Saxnäs, 29.VI.47
leg. Ringdahl. 9, Sodermanland, Groningen, 14.VI.54. 9, Angermanland, Docksta, 18.VI.54.
Q, Raa 5.V.46. 3 &, Skane, Väderön, 7.VII.34, 24.VIII.49 & 30.V.52. 4, Uppland,
Österskär, 12.VIII.48, 4, Arild, 11.11.25. 9, Vallbo, 29.VI.35. 9, Skane Bästad, 30.V.52.
4, Mölle, 19.V11.56, leg. Ringdahl. 4, Skane, Tyringe, 30.VIII.47. 6 9, Skane, Kullaborg
20.VIII.49. 9, Bolmen, 15.VI.54. & 2, Narke Mulhytton tf. Lillän, 5.VIII.52. 3 & (with
one preparation), ®, Närke, Hjälmarsberg, 6.VIII.52. 9, Gotland, Västerheide, 27.VI.50.
6, Angermanland, Hampnäs, 18.VI.54. 4, (with preparation), 2, Norbotten, Vagamo, 14.
VII.53. 4 & (with one preparation), Torne Lappmark, Björkliden, 12 & 17.VII.54, 2 8,
Torne Lappmark, Abisko-omr., 22—23.VII.54.
There are also accurate records for Scotland (SPENCER 1956) and France
(SéGuy 1934).
Distribution. Great Britain (incl. Scotland), France, Switzerland, Hol-
land, Sweden (including Lappland), Denmark, Germany, Austria, Czechoslovakia,
Hungary, Finland.
Several of the distribution records given by HENDEL (1931) were based on
misidentifications and must be deleted. In particular the Syrian specimen is A.
hiemalis Becker.
American records of this species (see FRICK 1959) need confirmation. FRICK
describes his nigripes as having a white squamal fringe, which is not the case in
European nzgripes.
The species is one of the commonest Agromyzids to appear in the sweep-net in
Northern Europe.
Host plants. Gramineae: tribes Glycerieae, Aveneae and Agrosteae. The
species is always in abundance on G/yceria maxima (Hartm.) Holmb. At the Ash
Vale locality I also observed it on G/ycerza fluitans (L.) R.Br. DE MEIJERE (1925)
also took this species on Holcus mollis L.
HERING (1957) keys this species under several other genera: Phalaris (3668),
Setaria (4811), Agropyron (151), Alopecurus (273) and Hordeum (2607).
Of these the Phalaris record is almost certainly incorrect: according to my ob-
servation in this country „igripes occurs freely on Glyceria but never on Phalaris
when the plants grow together. Some of the other records are probably correct but
they cannot be accepted without confirmation as they are not based upon bred
material, and considerable confusion has hitherto reigned over the larval characters
of this group. The following list of known food-plants will no doubt be enlarged
in the future, considering the ease with which ngripes is swept.
Normal food-plants: Glycerieae; Glyceria maxima (Hartm.) Holmb.; G. flustans
(L.) R.Br. Aveneae: Holcus (recorded from H. lanatus L. and H. mollis L.)
Recorded as food-plants once only: Aveneae: Deschampsia caespitosa (L.) Beauv.;
Agrosteae: Agrostis canina L. Old records of this species on Phragmites refer to
hendeli (described below).
One form of nigripes appears to merit subspecific rank.
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G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 133
Agromyza nigripes Meigen brachypodii subspec. nov.
Agromyza albipennis Buhr (nec Meigen), 1941. Stettin. ent. Ztg., vol. 102, p. 90. Syn. nov.
Textfigure 6a
Identical on external characters with small specimens of the typical form.
Squamal fringe ochreous. Small size characteristic; wing-length & 1.8 mm, 2
2.2 mm.
g genitalia. Aedeagal apodeme, 9th sternite, postgonites, pregonites and
aedeagal hood as for the typical form. Basiphallus consisting of two broad but
fairly ill-defined rods, with small ventral appendages (as in the typical form).
Mesophallus poorly sclerotised except at its outer edges. Distiphallus fairly small,
about as long as the basiphallus.
The ejaculatory bulb (Fig. 6a) is small, evenly rounded, transparent except for
a contrasting black patch on one side near its base. Stem strongly sclerotised and
fairly long.
Larva. Unknown.
Biology. BUHR (1941) states that puparia were found in a large communal
blotch-mine on 7.1X.30. Pupation is thus in the mine (in contrast with the
typical form). The mine specimens are unfortunately not extant.
Material examined. In the HERING collection. Type g' (with pre-
paration) Sagone, Corsica ex Brachypodium sylvaticum, leg. Buhr: emerged
17.IX.30. Allotype 9 same data, emerged 15.IX.30. Paratype g° (with prepa-
ration) same data, emerged 16.IX.30.
I erect this subspecies on account of the distinct ejaculatory bulb (not paralleled
in 38 preparations of the typical form from numerous localities) and the unusual
food-plant. Brachypodium is an easily recognised grass, and can be said witb
reasonable certainty not to be attacked by Agromyza spp. in this country. The
combination of these two facts with the unusual locality makes it probable that we
are dealing with a case of geographical variation. It will be interesting if further
material can be obtained from the Mediterranean region.
Distribution Corsica:
Host plant. Brachypodium sylvaticum (Huds.) Beauv.
Agromyza lucida Hendel 1920
Agromyza lucida Hendel, 1920, Arch. Naturgesch., vol. 84, part A, fasc. 7, p. 121.
(as subspecies of Agromyza reptans Fallen (1823)) (nec Hendel, 1922. Wien. ent. Ztg.,
vol. 39, p. 66, et 1931. Flieg. pal. Reg., vol. 59, p. 129: nec passim apud Hering, de
Meijere et alibi).
Agromyza holosericea Strobl (nec Bouché) 1893 Wien. ent. Ztg., vol. 12, p. 135, et
alibi. Syn. nov.
Agromyza airae Karl, 1926, Stettin. ent. Ztg., vol. 87, p. 138: etiam Hering, de Meijere
et passim.
Agromyza nigripes Stary (nec Meigen), 1930, pro parte. Acta Soc. Sci. nat. Morav., vol. 6,
p. 143. Syn. nov.
134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
Textfigures 7a, 7b, 7c, 18, 29, 40a, 40b, 40c, 40d, 50.
Agr. holosericea Strobl. (nec Bouché) is synonymised as a result of my examin-
ation of the specimen in HENDEL’s collection, thus determined in STROBL's hand-
writing. STARY's “nigripes” on Glyceria is clearly lucida as shown by his figure
of the larval mouthparts.
The type of A. lucida Hendel is mounted on a long fragile pin rendering
examination hazardous. It is a freshly emerged specimen whose colours are not
properly developed (as noted by HENDEL, 1931). Its specific identity is however
quite clear. The name lucida cannot be retained for the Phragmites-feeding species,
which is redescribed below as hendeli. The type was obviously not used for
HENDEL's description in 1931 (naturally enough in view of the fragile mount),
and the description given applies to the species described below as hendeli, not
the original /zcida. Consequently most subsequent authors have applied the name
lucida to the wrong species.
Frons usually broad, though sometimes approaching the narrow type, 1.0—1.5
times the width of an eye at the level of the front ocellus: parallel-sided or nar-
rowing forwards. Antennae usually clearly separated at their bases. In profile (Fig.
18) the orbits narrowly visible, the cheeks usually not so. Eyes large, slightly
pubescent. Jowls narrow: ratio of the eye-height to the height of the jowls 5—13.
Facial keel only slightly protruding in profile. 4 or 5 strong orbital bristles: lower
orbitals directed either backwards or inwards. 3rd antennal segment small and
rounded: antennal pubescence rather coarse. Frons dark-brown, sometimes yellow-
brown centrally. First two antennal segments usually brown or yellow-brown.
Acrostichals in 8—9 rows. Legs of variable colour: usually at least the front legs
with yellow tips to the femora, brown tibiae and yellowish tarsi. Sometimes also
the other tarsi brown and pale tips to all the femora.
Ratio of 2nd and 3rd costal segments (Fig. 29) 1.6—3; 2nd and 4th 2.7—4.
Wing-tip usually nearer the end of r4,, than m, ı 9. Wing-veins brown: mem-
brane clear or slightly brownish tinged. Squama white, grey or ochreous, usually
ochreous on its margin, with a thick black or dark brown fringe. Wing-length
22275 ORE
One of the largest species in the group.
3 genitalia. Distance from the tip of the aedeagal apodeme to the end of the
aedeagal hood 1 mm. Aedeagal apodeme more or less straight, the strengthened
part posterior to the attachment of the inner process of the postgonites occupying
rather more than 1/5 of the whole apodeme. 9th sternite fairly slender, somewhat
less rounded than in nigripes (compare Fig. 2), strongly bent downwards towards
its tip: short hypandrial apodeme, its tip reaching almost 2/3 of the length of the
aedeagal apodeme. Pregonites smaller than in zigripes (compare Fig. 2): post-
gonites a little larger, more quadrate. The aedeagal hood projects further forward
than that of albipennis (compare Fig. 1), and is fairly weakly sclerotised. Phallo-
phore as figured for albipennis (Fig. 1) but relatively longer.
Basiphallus (Fig. 7c) consisting of two short, fairly broad rods, the right being
a little broader than the left. A pale wedge-shaped sclerite lies in the membrane
beneath the basiphallus. Mesophallus consisting of two large elliptic plates, which
almost meet beneath the distiphallus. Distiphallus (Fig. 7b) fairly long, being
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 135
about as long as the basiphallus or clearly longer than it. Tensile membrane across
the top of the distiphallus slightly sclerotised.
Ejaculatory bulb (Fig. 7a) large and strongly sclerotised, often almost completely
black: about as broad as long. Stem short and curved, also strongly sclerotised.
Larva. Antennae small: maxillary palpus large and thick: slender but strong
longitudinal sclerite. Narrow mandibular abductor apodeme present. Mandibles
(Fig. 40a) long, with three teeth, the lower one being smaller than the upper
two: usually a slight protuberance posteriorly below the point of attachment of the
labial sclerite. 2—3 pairs of filaments above the mandibles.
Labial sclerite (Fig. 40b) long, its lower edge curved: paraclypeal phragma
more strongly sclerotised basally than apically; the ventral process is heavily
sclerotised for half its length: antero-dorsal process weak.
Head without spines. Prothoracic girdle consisting of about 10 rows of minute
spines: stronger ventrally. Mesothoracic girdle absent. Metathoracic girdle with
about 10 rows of small spines laterally; about 2 rows dorsally and ventrally.
Abdomen with more numerous spines than in other species: these are small but
sharply pointed: rather irregular but mostly pointing dorsally or postero-dorsally.
All bands at their broadest laterally, weaker ventrally: dorsal edge almost bare.
Ist segment with 16—20 rows of spines: 2nd to 5th segments with 14—16 rows:
6th with 10—12 rows: 7th with 7—8 rows (laterally only): 8th with 5—6 weak
rows (laterally only). Also numerous spines in the anal region.
Front spiracles (Fig. 40c) small, on a short twisting atrium, with 12—14 small
bulbs: the anterior horn slightly larger than the posterior. Hind spiracles (Fig.
40d) separated by 11/5—2 times their own width; their bulbs elongate-oval, rather
widely separated from each other: adjacent bulbs forming an angle of 90°. Atrium
similar to that of the front spiracles.
Length (full-grown) 4.5—5.5 mm.
Biology. Mine (Fig. 50) on upperside of leaf. Eggs laid singly on the edge
of the leaf, usually near the leaf-tip. The larva at first eats towards the leaf-tip,
but soon turns and produces a broad shallow blotch, on Deschampsia occupying
the whole leaf-breadth. Frass particles fine and inconspicuous. The mine is shallow
and the lower parenchyma is left untouched (contrast, nigripes). The larva crosses
the midrib freely. On Glyceria several eggs are often laid on one leaf and the
resulting mines coalesce to consume the whole leaf.
Pupation follows outside the mine, the puparia adhering to the leaf near the
mine. Puparia black or dark red.
Two generations. Flies occur in June and again in August. First generation
larvae found in June/July; second generation larvae in September/October (later
than „igripes).
Material examined. In the HENDEL collection. Type ®, Ost. sup., Altsee
26.VIII, leg. Czerny (swept). @ Stolp i. P., 10.11.26, ex Deschampsia caespitosa leg. Karl.
2 Kartashevka, Petrograd 2.VII.26, leg. Stackelberg. & (with preparation) 9, Alte Samm-
lung, Klostenenburg. ® "Agr. carbonaria Zett. B holosericea Bouché’ det. Strobl. 9, no
data (presumably Vienna region).
136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
In the author's collection. 3 4 (with 1 preparation), 4 2 Ash Vale, Surrey, 5.1X.60, ex
Glyceria maxima: emerged Oct. 60 and March/April 61. 4 larval preparations, Ash Vale,
Surrey, 5.IX.60 ex Glyceria maxima.
In the GROSCHKE collection. & (with preparation) Soritz, Beutzen, 28.VII.27, ex
Deschampsia caespitosa leg. Schülze. 9 Mecklenburg, Rostock, 16.VII.49, ex Deschampsia
caespitosa, no. 462, Buhr.
In the HERING collection. & paratype of the synonymous a/rae Karl (with preparation)
ex Deschampsia caespitosa. 9 Chippenham Fen, Cambs., emerged 22.VII.58, ex Deschampsia
caespitosa, Hering, 6317. 2 © Soritz, Beutzen 17.VII.27, ex Deschampsia caespitosa, leg.
Schütze. @ Mecklenburg, Rostock, ex Deschampsia caespitosa, no. 479, Buhr. & (with
preparation), Mecklenburg, Rostock, 25.VII.31, ex Echinaria capitata, leg. Buhr. © Crossen-
an-Oder, 7.V.30, ex Glyceria maxima Hering, 3598. ® swept at Stempeda, Harz, 25.V1.27,
Hering, 3179.
In the NowakowskI collection (Warsaw). @ 16.1.16, ex Deschampsia caespitosa, leg.
Karl.
In the RYéN collection (Hälsinborg, Sweden). 2 9 Västergötland, Ulricehamn, 27.VI.47.
& Docksta, Angermanland.
In addition to these records DE MEIJERE (1943) knew this species from Hol-
land, and Stary figures the larva from Czechoslovakia.
Distribution. England, Holland, Sweden, Germany, Austria, Czechoslova-
kia, Poland, Russia. Not uncommon where its host-plants grow: turns up sparingly
in swept material.
Host plants. Gramineae: Tribes Glycerieae, Festuceae and Aveneae. Nor-
mal food-plants: Glycerieae; Glyceria maxima (Hartm.) Holmb. Aveneae:
Dschampsia caespitosa (L.) Reauv. Recorded as food-plant once only: Festuceae:
Echinaria capitata.
HERING (1957) also keys this species under Agropyron (150), Bromus (955)
and Holcus (2588). I regard these records as requiring confirmation, as they are
not based upon bred material.
Agromyza hendeli sp. nov.
Agromyza nigripes Kaltenbach (nec Meigen), 1856, Verh. naturh. Ver. preuss. Rheinl. &
Westph., vol. 13, p. 242, et 1874, Die Pflanzenfeinde aus der Klasse der Insekten, p. 764:
etiam de Meijere pro parte, Hering pro parte et passim.
Agromyza lucida Hendel 1922 (nec Hendel 1920), Wien. ent. Ztg., vol. 39, p. 66, et
1931, Flieg. pal. Reg., vol. 59, p. 129: etiam passim apud Hering, de Meijere et alibi.
Textfigures 8a(i), 8a(ii), 8b, 19, 30, 41a, 41b, 41c, 41d
Examination of the type of lucida shows that the name can no longer be used
for this species. A new name is required although the species has been known for
over a hundred years.
Frons narrow, 0.8—1.2 times the width of an eye at the level of the front
ocellus; narrowing somewhat forwards. Antennae practically touching at their bases.
In profile (Fig. 19) the cheeks usually not visible except very narrowly beneath
the eye. Orbits barely visible. Eyes large, faintly pubescent; their height 4—8
times the height of the jowls. 5—6 strong orbital bristles, all directed more or
less posteriorly: no clear distinction between upper and lower orbitals. Third
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 137
10b
Figs. 8—10. Male genitalia of Agromyza: 8, hendeli sp. nov.; 9, spenceri sp. nov.; 10,
bhragmitidis Hendel: a — ejaculatory bulb; b — lateral view of phallus or distiphallus;
c — ventral view of phallus
138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
antennal segment large, broadened apically, somewhat deeper than in zigripes.
The row of hairs on the second segment strongly developed. Peristomal hairs
often longer and more numerous than usual; bristles at the end of the palp strongly
developed. Usually the 1st and 2nd antennal segments and the frons are brown,
though sometimes the head is completely black.
Acrostichals in 6—9 rows. Legs with usually at least the front femora yellow
at their tips; sometimes the tips of all femora yellow and the front tarsi brown.
Ratio of 2nd and 3rd costal segments (Fig. 30) 2.5—3; 2nd and 4th 3.5—4.8.
Wing-tip between the ends of r, , 5 and m, + 4 or nearer the latter. Veins usually
pale (compare albipennis): membrane clear. Squama white, with a contrasting
black or dark brown fringe. Wing-length 2.0—2.6 mm.
g genitalia. Distance from the tip of the aedeagal apodeme to the end of the
aedeagal hood about 1 mm. Aedeagal apodeme almost straight, strongly strength-
ened posterior to the attachment of the inner processes of the postgonites, this
part occupying 1/5 of the whole apodeme. 9th sternite small, rather rounded, with
a well-defined hypandrial apodeme occupying 1/5 of the total length of the 9th
sternite: its tip reaches only to 1/5 of the length of the aedeagal apodeme.
Pregonites smaller than those figured for nigripes (Fig. 2): postgonites more
squarely shaped. Aedeagal hood very large, but weakly sclerotised, in profile
clearly raised above the level of the aedeagal apodeme. Phallophore less indented
posteriorly when viewed in profile than that figured for albipennis (Fig. 1).
Basiphallus consisting of two broad rods, dentate on their lower edge with
additional appendages ventrally (Fig. 8b). These are larger than in other species
(e.g., nigripes) and at least partially composed of dark sclerotin: particularly the
posterior edge is clearly defined. Mesophallus consisting of a pair of ill-defined
lobes, strongly sclerotised on their outer edges (in profile this gives the impression
of a second pair of appendages from the basiphallus). Distiphallus with its rear
part strongly bent downwards; also the tip obviously bent down (more markedly
so than in nigripes). Distiphallus about equal in length to the basiphallus.
Ejaculatory bulb (Figs. 8a(i), Sa(ii) strongly sclerotised, appearing black or
dark brown; very narrow, at least twice as long as broad. Stem long, more or less
straight.
Larva. Antennae small; maxillary palpus fairly large: slender longitudinal
sclerite. Narrow mandibular abductor apodeme present. Mandibles (Fig. 41a) large
and powerful with two teeth. Smal mandibular adductor apodeme present.
2—3 pairs of inconspicuous filaments above the mandibles.
Labial sclerite (Fig. 41b) somewhat curved, especially on its lower edge: para-
clypeal phragma with its processes composed of brown and black sclerotin (contrast
phragmitidis): antero-dorsal process very small.
Head with a few small scattered spines below the mandibles, otherwise bare.
Prothoracic girdle consisting of about 8—10 rows of minute spines; mesothoracic
girdle absent, or at most a few spines in two rows dorsally. Metathorax and first
five abdominal segments with strong bands of large spines; in each band the
posterior 3 or 4 rows consist of larger spines pointing more or less posteriorly; the
other spines point generally dorsally or postero-dorsally; bands weakening dorsally
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 139
and ventrally, the dorsal edge of the abdomen being quite bare, the ventral nearly
so. Metathorax with about 8 rows of spines; 1st to 4th abdominal segments with
10—14 rows; Sth with about 8 rows; 6th to 8th segments with some scattered
rather blunt spines laterally in 3—4 rows, but these bands very inconspicuous by
comparison with those of the preceding segments. Also the anal region with some
spines.
Front spiracles (Fig. 41c) small, with two almost equal horns bearing 10—12
small bulbs: atrium relatively short, evenly curved but not twisted. Hind spiracles
(Fig. 41d) separated by their own width or a little more: their bulbs elongate-oval,
almost touching each other centrally; adjacent bulbs forming an angle of 60°.
Length (full-grown) 3.75—4.5 mm.
Biology. Mine (compare Fig. 51) on upperside of leaf. Eggs laid usually
in a row of 3 or 4. The resulting larvae produce a communal mine, at first eating
towards the leaf-tip in a gradually broadening communal channel. In the third
instar the larvae often split up producing a large irregular blotch, often too there
are several egg-groups on the same leaf, and the resulting mines coalesce, Frass
in large scattered particles; zig-zag feeding lines often visible in fresh mines.
Pupation follows normally outside the mine, the puparia adhering to the leaf
near the mine. Puparia black or brown.
Two generations. The larvae have been found in June and August: the second
generation is earlier than that of phragmitidis. Flies have been swept from June
to August.
Material examined. In the HENDEL collection. Type 4 with preparation),
Grünwald, emerged 21.11.20, ex Phragmites communis, leg. Hering, 1331. Allotype 9,
Bredow b. Nauen, emerged 9.1V.23 ex Phragmites communis, leg. Hering, 2154. Paratypes
Q, Wien, 19.VII.78, leg. Mik. 4, no. data leg. Bgst.
In the author’s collection. Paratypes. 2 & (with preparations), swept at Woodwalton Fen,
Hunts., 21 & 25.VIII.60. 2 9, Woodwalton Fen, Hunts., 16.VIII.60, ex Phragmites com-
munis: emerged 12.IX.60. 5 larval preparations from Woodwalton Fen, Hunts., 10.VIII.60,
22.VIII.60 and 21.VI.61.
In K. A. SPENCER’s collection. Paratype & (with preparation), Lake Garda, Italy, 25.X.56,
ex Phragmites communis: emerged 10.11.57.
In the GROSCHKE collection. Paratypes: ¢, Bredow b. Nasar, 12.1V.23, ex Pragmites
communis, Hering, No. 2154. 4 9, Baienfurt, Wttbg. (Fuchsenloch), 17.V1.55, leg.
Groschke. 4, Mecklenburg, Mönkweden, 1.IV.50, ex Phragmites communis, Buhr 600/49.
In the HERING collection. Paratypes: 4 (with preparation), Soritz, Beutzen, 15.VIII.27,
ex Phragmites communis, leg. Schütze. Q, Soritz, Beutzen Spring, ‘28, ex Phragmites com-
munis, leg. Schütze. 2 9, Berlin, Botanische Garten, 16.VII, ex Phargmites communis, no.
58, Hering 1951. ® (with preparation), Grünewald, April 1930, ex Phragmites communis,
1331, Hering. Also a larval preparation ex Phragmites communis, Berlin, Bot. Gart., 18.VI.
Sy
In the NowaKowsKI collection. Paratype ¢, Puzcza Kampinoska, Mfokiny, 29.VI.57;
puparium 1.VII, imago 11.VII, ex Phragmites communis, Caricetum, leg. Nowakowski, Inst.
Zool. P.A.N., Warszawa, No. 20/57.
In the RyDéN collection. Paratypes: 3 & (with 1 preparation), 2 9, Skane, Landskrone,
7.VI.50 and 15.VIII.52.
(22 paratypes in all.)
140 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 141
In addition to the above records DE MEIJERE knew this species from Holland
and SPENCER (1956) records it (as lucida) from Scotland. There are many other
references to the mines of this species having been observed (as nigripes and
lucida): however only records resulting from bred material can be trusted as the
mines of hendeli, spenceri (described below) and phragmitidis are not disting-
uishable, and the larval characters have been hitherto confused.
Distribution. Great Britain (incl. Scotland), Holland, Sweden, Germany,
Austria, North Italy, Poland. Common in northern Europe where its host occurs.
Host plant: Phragmites communis Trin.
Agromyza spenceri sp. nov.
Textfigures 9a, 9b, 20, 31, 43a, 43b, 43d
Frons narrow, 1—1.2 times the width of an eye at the level of the front ocellus,
strongly narrowing forwards. Antennae practically touching at their bases. In
profile (Fig. 20) the cheeks not visible, the orbits barely so. Eyes large, faintly
pubescent, their height 6—8 times the height of the jowls. Face-line very flattened
in profile. 4 strong orbital bristles, all directed more or less backwards. Third
antennal segment small and rounded (compare phragmitidis). Frons and first two
antennal segments brown: head otherwise black.
Acrostichals in 8 rows. Legs with the front femora broadly yellow at their tips,
the others narrowly so. Front tarsi yellow. Legs otherwise dark brown.
Ratio of 2nd and 3rd costal segments (Fig. 31) 2.5—3: 2nd and 4th 4—4.2
(contrast lucida). Wing-tip between the ends of r, + „ and m,4 9. Wing veins
brown: membrane brownish tinged. Squama white with an ochreous margin and
strong jet-black fringe. Wing-length 2.2—2.6 mm.
3 genitalia. Distance from the end of the aedeagal apodeme to the end of the
aedeagal hood 0.8 mm. Aedeagal apodeme stout, almost straight: the strengthened
part posterior to the attachment of the inner processes of the postgonites occupying
1/5 of the whole. 9th sternite stout, broad and rounded, bearing a well-defined
hypandrial apodeme: this occupies about 1/, of the total length of the 9th sternite:
its tip reaches 3/5 of the length of the aedeagal apodeme. Pregonites large, as
figured for nigripes (Fig. 2), but forming a rounded lobe on their inner anterior
corners. Postgonites as figured for nigripes (Fig. 2). Aedeagal hood strongly
sclerotised at its edges. Phallophore as shown for albipennis (Fig. 1).
The sclerotisation of the basiphallus and mesophallus is weak and ill-defined.
Basiphallus short and slender, though its right rod slightly broader than its left.
Mesophallus very faintly sclerotised except on its outer edge, forming a large ill-
defined lobe beneath the distiphallus. The distiphallus (Fig. 9b) is very large and
of characteristic shape, being elongated in its central part: the membrane stretching
across the top of the distiphallus is partially sclerotised along its edge. Distiphallus
about 11/5 times as long as the basiphallus.
Figs. 11—13. Male genitalia of Agromyza: 11. prespana Spencer; 12, graminicola Hendel;
13, baetica sp. nov. a — ejaculatory bulb; b — lateral view of phallus or distiphallus;
c — ventral view of phallus; d — aedeagal apodeme and 9th sternite in ventral view
142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
Ejaculatory bulb (Fig. 9a) narrow, moderately sclerotised in its basal 2/, more
weakly so in its distal 1/3: faint medial streak visible. Stem straight.
Larva. There are no extant larval specimens. However the following charac-
ters have been described from the puparia. Mandibles (Fig. 43a) with 2 strong
teeth (as in hendel: and phragmitidis). Labial sclerite (Fig. 43b) very long:
processes of the paraclypeal phragma consisting of brown sclerotin. Hind spiracles
(Fig. 43d) separated by a little over thelr own width: adjacent bulbs forming an
angle of 70°.
As far as these characters go the larvae cannot yet be distinguished from those
of hendell.
Biology. The mines are identical with those of hendeli and phragmitidis
(compare Fig. 51): the description under those species applies here also.
The puparia are reddish brown and are attached to the leaf near the mine.
Probably only one generation, as larvae taken in late July did not produce flies
until the following year.
Material examined.
In K. A. SPENCER’s collection. Type & (with preparation) collected on 26.VII.
54, at a small lake between Briancon and Argentières, Basses Alpes, France, at an
elevation of about 3,100 ft. (946 metres): ex Phragmites communis; emerged
26.VI.55. Paratypes: 2 J and 3 9, same data, emerged 23—29.IV.55.
Distribution. French Alps.
Host plant. Phragmites communis Trin.
Figs. 14—15. Male genitalia of Agromyza. 14, distorta Griffiths; 15, oryzae (Munakata).
a — ejaculatory bulb; b — lateral view of phallus; c — ventral view of phallus
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 143
Agromyza phragmitidis Hendel 1922
Agromyza phragmitidis Hendel, 1922, Wien. ent. Ztg., vol. 39, p. 65.
Textfigures 10a, 10b, 10c, 21, 32, 42a, 42b, 42c, 42d, 51
Frons narrow, 0.8—1.2 times the width of an eye at the level of the front
ocellus: narrowing forwards: the antennae practically touching at their bases. In
profile (Fig. 21) the cheeks not visible, and the orbits not or barely so. Eyes very
large, slightly pubescent. Jowls normally extremely narrow: ratio of eye-height
to the height of the jowls 5—11 (in one specimen the jowls are almost elimin-
ated). 5 weak orbital bristles, normally clearly divided into 2 upper orbitals and
3 lower, the latter nearer to each other than the former: often the orbitals are
rather small and adpressed to the frons. 3rd antennal segment small and rounded.
Frons, orbits and first two antennal segments clearly brown, sometimes yellow-
brown. Arista yellow basally.
Acrostichals 8-rowed. Legs with the tips of all the femora clearly yellow, the
front femora more broadly so. All tarsi at least brown, often clearly yellow.
Ratio of 2nd and 3rd costal segments (Fig. 32) 2.5—3.2; 2nd and 4th 3.5—4.5.
Wing-tip just after the end of ry 45. Costa often weak between r, 4, and mj 4 9.
Figs. 16—19. Heads in profile of Agromyza: 16, albipennis Meigen; 17, nigripes Meigen;
18, lucida Hendel; 19, hendeli sp. nov.
144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
Wing-veins pale: membrane clear or whitish. Squama white with a white fringe.
Wing-length 2.2—2.8 mm.
A large species. The basic colour of the whole body is sometimes dark brown,
not black.
3 genitalia. Distance from the end of the aedeagal apodeme to the end of the
aedeagal hood a little over 1 mm. Aedeagal apodeme more or less straight: the
strengthened part posterior to the attachment of the inner processes of the post-
gonites occupying nearly 1/, of the whole apodeme. 9th sternite long and narrow,
more or less straight-sided in dorsal view; in lateral view obviously bent down-
wards towards its tip. Hypandrial apodeme large, but not clearly differentiated,
being continuous with the rest of the 9th sternite: it occupies about 1/3 of the
total length of the sternite: its tip reaches 2/3 of the length of the aedeagal
apodeme. Pregonites much smaller than those figured for nigripes (Fig. 2):
postgonites as figured. Aedeagal hood large, weakly sclerotised, clearly raised
above the level of the aedeagal apodeme in profile (compare hendelt). Phallo-
phore forming a short double ridge in dorsal view, lacking the indentation shown
posteriorly in albipennis (Fig. 1).
Basiphallus (Fig. 10c) consisting of two long clearly defined rods of moderate
width. Mesophallus consisting of two triangular plates which draw together
beneath the distiphallus. Distiphallus (Fig. 10b) large and thick: its rear part bent
downwards: a little longer than the basiphallus.
Ejaculatory bulb (Fig. 10a) narrow (but not so narrow as in hendel), strongly
sclerotised, appearing black or brown. Its stem long, somewhat curved.
Larva. Antennae small: maxillary palpus large, long: longitudinal sclerite
strongly developed. Mandibular abductor apodeme weak, barely visible. Mandibles
(Fig. 42a) large and powerful with two teeth. Sometimes a small mandibular
adductor apodeme present. 3 pairs of long filaments above the mandibles: these
clearly reaching beyond the mandibular teeth.
Labial sclerite (Fig. 42b) long, slightly curved: paraclypeal phragma strongly
sclerotised only at its base: all three processes consisting of transparent sclerotin:
the ventral process is open at its tip: end of the dorsal process clearly separated
from the medial process: antero-dorsal process vestigial.
Head bare apart from 2 or 3 small spines beneath the mandibles. Prothoracic
girdle consisting of about 12 rows of minute spines. Mesothoracic girdle absent,
or only a small patch of spines present ventrally. Metathorax and first four
abdominal segments with strong bands of large spines: in each band the posterior
3—4 rows consist of larger spines pointing more or less posteriorly: the other
spines pointing dorsally or posterodorsally. Bands weakening dorsally and ventral-
ly: the dorsal edge of the abdomen almost completely bare: only the bands of
the first two abdominal segments continue over the ventral edge. Metathorax
with 8—10 rows of spines: Ist and 2nd abdominal segments with 10—12: 3rd
segment with 8—10 rows: 4th with 6—8 rows: 5th segment with a very weak
band of spines present only laterally in about 4 rows: 6th and 7th segment without
bands: 8th segment band represented by a few spines ventrally: also a few spines
in the anal region.
Front spiracles (Fig. 42c) fairly small, two-horned, the anterior horn being
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 145
slightly longer than the posterior: they bear 10—12 bulbs: atrium somewhat
twisted. Hind spiracles (Fig. 42d) separated by about their own width: their
bulbs elongate-oval, almost touching each other centrally: adjacent bulbs forming
an angle of 60°—70°.
Length (full-grown) 4.5—6 mm.
Biology. The mines (Fig. 51) are identical with those of hendeli; the
description given under that species applies here also. Pupation follows normally
outside the mine: the puparia adhere to the leaf near the mine. Puparia very
variable in colour, ranging from black to almost yellow.
Two generations. The larvae are particularly common in late September and
October. Adults have been swept in May/June and again in September. The
autumn generation is later than that of hendels.
Material examined. In the HENDEL collection. Types 4 9 (both on the same
mount), Güntersberg-an-Oder, 6.III.29, ex Phragmites communis, leg. Hering.
2 6,2 9, same data, emerged Feb-Apr. 29, Hering, no. 729. 9, Verebély, Kertész, May
33. ®, Austria, Alte Sammlung, labelled ‘’carbonaria det. Schiner”.
In the author's collection. & (with preparation), Woodwalton Fen, Hunts., 23.VIII.60, ex
Phragmites communis: emerged 27.X.60. 69, same data, emerged April 1961.
Larval preparations ex Woodwalton Fen, Hunts., ex Phragmites communis.
In K. A. SPENCER's collection. 2 ¢ (with preparations), Wonwells, Devon, 15.IX.54, ex
Phragmites communis: emerged 26.V.55 and 1.VI.55.
22a
Figs. 20—23. Heads in profile of Agromyza: 20, spenceri sp. nov; 21, phragmitidis Hendel;
22, ® prespana Hendel; 22a, male antenna of same; 23, graminicola Hendel
146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
In the GROSCHKE collection. & 9, Güntersberg-an-Oder, 28.11.29, ex Phragmites communis,
leg. Hering. 4, Neü-Freier (?), 29.IX.53, ex Phragmites communis.
In the HERING collection. 3 & (with 1 preparation), 9, Güntersberg-an-Oder, ex Phrag-
mites communis: emerged Feb. '29: Hering, 3348. 4 (with preparation), 9, Crossen-an-
Oder, Sept., ex Phragmites communis. & (with preparation), Soritz, Beutzen, ex Phragmites
communis, Spring, 1928, no. 62, leg. Schütze. 9, same data, emerged 29.IX.27, leg. Schütze.
Puparial preparations from Bradons b. Nassau, end Sept, 1922: and St. Peter-Bord,
10.VIII.54, ex Phragmites communis, leg. Ludwig.
In the NowakowskI collection. & (with preparation), Puzcza Kampinoska, Mfokiny, ex
Phragmites communis; Saliceto-Populetum, larva 15.IX.55, puparium 27.IX, imago 11.V1.56,
leg. Nowakowski: Inst. Zool. P.A.N., Warszawa, 60/57.
In the RYDÉN collection. 9, Raa, Skane, 2.VIII.35.
Besides the above records DE MEIJERE (1925) knew this species in Holland
(as nigripes), and SASAKAWA (1956) records it for Japan. There are some other
published records based on the mines being observed, but these require confirm-
ation, as the mines are not distinguishable from those of spenceri and hendeli.
Distribution. England, Holland, Sweden, Germany, Austria, Hungary,
Poland, Japan. Common in northern Europe where its host grows.
Host plant. Phragmites communis Trin.
Agromyza prespana Spencer 1957
Agromyza prespana Spencer, 1957, Ent. mon. Mag., vol. 93, p. 35.
Agromyza sp. b Hering, 1953, Tijdschr. PIZiekt., vol. 59, p. 109—191.
Textfigures 11a, 11b, 11c, 22, 22a, 33, 44a, 44d
Frons 1.4—1.5 times the width of an eye at the level of the front ocellus,
25
Figs. 24—26. Heads in profile of Agromyza: 24, baetica sp. nov.; 25, distorta Griffiths;
26, oryzae (Munakata)
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 147
slightly narrowing forwards; the antennae are separated at their bases. In profile
(Fig. 22) the eye is large, its height about 7 times the height of the jowls at
their centre. The orbits project above the antennae. Cheeks extremely narrow,
almost eliminated below the antennae. 2nd antennal segment with its upper edge
lengthened, so that the 3rd segment is directed downwards. 3rd antennal segment
sexually dimorphic: in the male (Fig. 22a) larger with a very long fringe of
upturned hairs: in the female (Fig. 22) a little longer than high with the fringe
of hairs shorter. Facial keel clearly projecting in profile. 2 + 3 orbital bristles,
fairly weak, the lower 3 directed inwards. Ocellar bristle also weak. Frons dark
brown, first two antennal segments, sometimes also the third, yellow-brown, orbits
and ocellar triangle shining black, lunule grey.
Acrostichals in 6—8 rows. Mesonotum very shining black. Legs with the tips
of the femora yellow (especially the front pair) and all tarsi yellow-brown;
sometimes also tibiae yellow-brown. The pubescense of the legs is short and fine.
Ratio of 2nd and 3rd costal segments (Fig. 33) 2.8—3.2; 2nd and 4th 3.6—4.2.
Wing-tip nearer the end of ry ‚5 than m, +5. Veins yellow-brown: membrane
clear. Squama clear or white with a contrasting black fringe. Wing-length
19 2]ernm.
3 genitalia. Distance from the end of the aedeagal apodeme to the end of the
strengthened portion posterior to the attachment of the inner processes of the post-
gonites occupying slightly less than 1/; of the whole apodeme. Pregonites
small, but strongly sclerotised, with obvious sensory pores on their inner edges.
Postgonites small, club-shaped. Aedeagal hood very small and narrow, but
strongly sclerotised. Phallophore much as in albipennis (Fig. 1), but with its
postero-dorsal edge straight and strongly sclerotised.
Phallus fairly typical of the group. Basiphallus (Fig. 11c) consisting of two
short rods, the right being broader than the left. Mesophallus consisting of two
more or less triangular plates, pointed where they almost meet beneath the disti-
phallus (Fig. 11b). Distiphallus large, clearly longer than the basiphallus, its
rear part strongly bent downwards.
Large characteristic ejaculatory bulb (Fig. 11a), broadened out like a fan
distally. Stem by comparison very small.
Larva. The following characters have been described from the puparia.
Mandibles (Fig. 44a) very small and narrow, with two teeth. Small pro-
tuberance posteriorly beneath the point of attachment of the labial sclerite.
Hind spiracles (Fig. 44d) situated on large projections: separated by over three
times their own width: their adjacent bulbs forming an angle of about 90°.
The larval specimen figured by HERING (1953) as Agromyza sp. b” is almost
certainly identical with this species. This specimen shows a moderate-sized maxil-
lary palpus, small antenna and a long narrow longitudinal sclerite (as normally
in this group): also a clearly defined mandibular abductor apodeme. Labial
sclerite strong and straight: processes of the paraclypeal phragma strongly sclerot-
ised, but not widely splayed (contrast graminicola). Prothoracic girdle with about
6 rows of minute spines: mesothorax bare. Metathorax and first six abdominal
segments with strong bands of spines, mostly directed posterodorsally: all bands
148 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
broadest laterally, but strong dorsally too, continuing over the dorsal edge with
several rows of spines: weaker ventrally. All spines somewhat blunt. Metathorax
with about 10 rows laterally; first three abdominal segments with 14—18 rows;
4th and 5th with 10—14 rows; 6th with 12 rows of fine spines laterally, much
weaker dorsally and ventrally: 7th and 8th segments and the anal region nearly
bare. Front spiracles with 11—14 bulbs; bulbs of the hind spiracles character-
istically twisted in side view (see HERING’s figure).
Biology. There is one extant mine-specimen of Agr. spec. b. This is a
gradually broadening mine beginning at the leaf tip, length about 4 cm, maximum
width 4 mm. The frass particles are clearly defined.
Pupation probably outside the mine. The puparia are large and reddish.
One generation. Flies have been swept only in May/June. GROSCHKE's bred
material did not emerge until the following May.
Material examined. In the British Museum (Natural History). Type ¢, Lake
Prespa, Macedonia, swept from vegetation near the lake, mid-June 1955, leg. Coe, B.M.,
1955—460.
2 & (with one preparation), 3 9, paratypes, same data. 9, paratype, Lake Ochrid,
Macedonia, early June 55.
® Polje, Dalmatia, Novigrad (in valley amid olive trees etc.), 27—31.V.58, leg. Coe,
B.M., 1958—417.
In K. A. SPENCER’s collection. ¢ (with preparation), 9, paratypes; same data as type.
In the HENDEL collection. @ Prater, Wien, 19.V, leg. Hendel.
In the GROSCHKE collection. 9, Offingen, Wttbg., 24.V.55, ex Triticum, leg. Groschke.
®, Nekarrems, Wttbg. 18.V.55, ex Triticum, leg. Groschke. ® Berlin, Frohnen, 24.VI.28,
leg. Hering.
In the RyDpéN collection. 9, Skane, Kullaberg, 20.VI.48.
In the HERING collection (Berlin). Larval preparation of "Agromyza sp.b’, Görlitz,
Kunnersdorf, 20.VI.52, ex Triticum aestivum, Hering, no. 5860. |
Besides these records the puparia which DE MEIJERE (1937) described from
Czechoslovakia may belong here (see also HERING, 1957, no. 5305), but the
description is inadequate.
Distribution. Southern Sweden, Germany, Austria, Jugoslavia (Dalmatia
and Macedonia). It seems probable that this species was originally a southern
form, which has spread northwards by becoming associated with wheat. In the
Jugoslavian localities it is most unlikely that wheat was the food-plant.
Host plant. Triticum aestivum L.
Agromyza graminicola Hendel 1931
Agromyza graminicola Hendel, 1931, Flieg. pal. Reg., vol. 59, p. 122.
Dexthigures) 124, ODA ZR 23) 34, 452.,455,.45d.,52
Frons narrow, 1—1.3 times the width of an eye at the level of the front
Figs. 27—37. Wing maps of Agromyza: 27, albipennis Meigen; 28, nigripes Meigen; 29,
lucida Hendel; 30, hendeli sp. nov.; 31, spenceri sp. nov.; 32, phragmitidis Hendel;
33, prespana Spencer; 34, graminicola Hendel; 35, baetica sp. nov.; 36, oryzae (Munakata);
37, distorta Griffiths
] i 14
er } roup 9
visi Agromyza nigripes 14
SFITHS : Revision (0) 8
I mm
150 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
ocellus: usually narrowing forwards. Antennae narrowly separated at their bases.
In profile (Fig. 23) the cheeks are narrowly visible, but not clearly defined, below
the eye. Orbits narrowly visible. Ratio of the eye-height to the height of the jowls
4—9. Eyes bare. 4—5 orbital bristles, not clearly differentiated into upper and
lower orbitals: the third orbital directed more or less backwards: all orbitals often
fairly small and weak. 3rd antennal segment very large, its lower margin evenly
curved. Pubescence of antennae and arista fine. Frons and first two antennal seg-
ments dark brown.
Acrostichals 8-rowed. Legs dark brown to black, the front femora broadly
yellow at their tips: tarsi sometimes brown.
Ratio of 2nd and 3rd costal segments (Fig. 34) 2.7—3.6: 2nd and 4th usually
5—5.8 (rarely as little as 4): 4th segment usually very short. Wing-tip between
the ends of r4 5 and m, ı o. Veins pale: membrane clear. Squama white with
a white fringe. Wing-length g2.0—2.2 mm, 9 2.4—2.6 mm.
d' genitalia. Distance from the end of the aedeagal apodeme to the end of the
aedeagal hood 0.8 mm. Aedeagal apodeme thickened at its anterior end, more or
less straight. The strengthened part posterior to the attachment of the inner proces-
ses of the postgonites is relatively short, occupying 1/4 of the whole apodeme.
9th sternite stout and rounded, bearing a long hypandrial apodeme, which
occupies about 2/5 of the total length of the sternite: 9th sternite symmetrical
(contrast baetica): tip of the hypandrial apodeme reaching nearly 2/3 of the
length of the aedeagal apodeme. Pregonites large, as figured for nigripes (Fig. 2),
but with an anteriorly directed lobe on their inner anterior corners. Postgonites as
figured for nigripes (Fig. 2): inner processes somewhat thickened where they
curve down beneath the aedeagal hood. Aedeagal hood much deeper in profile
than that figured for albipennis (Fig. 1): it is strongly sclerotised on its posterior
surface. Phallophore relatively larger than that figured for albipennis (Fig. 1).
Basiphallus (Fig. 12c) short and broad, asymmetrical, the left rod being
obviously broader than the right. At the junction of the mesophallus and disti-
phallus on the left side a large downwards directed lobe. Mesophallus modified
to form two club-shaped appendages, which lie beneath the distiphallus: the left
appendage is slightly larger. Distiphallus (Fig. 12b) very large, a little longer than
the basiphallus: its rear part bent downwards.
Ejaculatory bulb (Fig. 12a) fairly broad with a ribbed appearance: a characer-
istic ’’waist’’ at its base (contrast baetica): its sclerotisation moderate. Stem short,
more or less straight.
Larva. There are no extant larval specimens. However the following
characters have been described from the puparia. Mandibles (Fig. 45a) with 2
strong teeth: not so broad as those of hendeli, spenceri and phragmitidis. Labial
sclerite (Fig. 45b) strongly thickened posteriorly: paraclypeal phragma strongly
sclerotised, its processes large and widely splayed. Hind spiracles (Fig. 45d)
separated by about 3 times their own width, with elongate-oval bulbs: adjacent
bulbs forming an angle of 90°—100°.
Biology. Mine (Fig. 52) on upperside of leaf. Eggs laid singly near the
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 15,
edge of the leaf. The resulting larva produces a gradually broadening channel at
first towards the leaf-tip, then turns towards the base of the leaf. Frass fairly
conspicuous, in two irregular rows. Width of the mine channel not exceeding
4 mm.
Pupation follows in the mine (HERING, 1951a); puparia reddish-brown.
HERING (195la) gives some interesting comments on the biology of this
species. According to his observations it never attacks Phragmites standing in
water, but only dwarf plants growing on quite dry ground.
At least two generations. Larvae taken in October produced flies in October
and the following year. RyDéN swept the flies in May and August.
Material examined. In the HENDEL collection. Type ¢ 2 (both on the same
mount), Crossen-an-Oder, emerged 10.X.34, ex Phragmites communis, Hering, no. 4295.
3 ©, labelled Alte Sammlung, Wien. 4, Hungaria, Lac. Fert., leg. Podkorny (with
genitalia extruded).
(The original type mentioned by HENDEL (1931) was bred in the Vienna region, and
must be presumed lost. There is no room for doubt as to the identity of this species.)
In the author's collection. 4 (with preparation), Crossen-an-Oder, emerged 9.III.35, ex,
Phragmites communis, Hering, no. 4295.
In the GROSCHKE collection. 9 ¢, same data as type.
In the HERING collection. & (with preparation), 2 9, same data as type, emerged Oct. 34
and March 35, Hering, no. 4295.
Also a puparial preparation ex Crossen-an-Oder, 8.X.34, ex Phragmites communis. Hering,
no. 4295.
In the NOWAKOWSKI collection. & (with preparation), Szeroky Bor, Poland, ex Phrag-
mites communis: puparium 17.VIII.58, imago 23.VIII, leg. Szepanski.
In the RyDéN collection. 9, Raa, 26.V.25. & (with preparation), Skane, Skälderviken,
15.VIII.55, leg. Ringdahl. 2 & Skane, Landskrone, 15 & 18.VIII.52.
o cS d on
= Oo (È gui Fe od o
E œ PA
d AG Oi On en
o ao er d Ga) Ce
œ
38e
38C
38a 38d
Fig. 38. Larval parts of Agromyza nigripes Meigen: a — mandibles; b — larval mouthparts;
c — front spiracles; d hind spiracles: e — section of the spinal band of the first
abdominal segment laterally
152 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
HERING (1951) also observed this species in the Stockholm area, so that its
range extends quite far north.
Distribution. Sweden, East Germany, Poland, Austria, Hungary.
Apparently this species does not reach Western Europe.
Host plant. Phragmites communis Trin.
HERING (1957) also keys this species under Arundo. I do not know the
origin of this record, but must regard it as requiring confirmation, especially in
view of the discovery of a related species in the Mediterranean area (described as
baetica below).
Agromyza baetica spec. nov.
Mextitoures 13a, Ml DB Reid ANSE
Frons fairly broad, 1.5 times the width of an eye at the level of the front
ocellus: parallel-sided. Antennae separated by the width of their first segment at
their bases. Cheeks and orbits clearly visible in profile (Fig. 24), especially the
orbits above the antennae. Ratio of the eye-height to the height of the jowls
4—7. Eyes with obvious pubescence. 5—6 weak orbital bristles, all directed more
or less backwards. Antennae with very large 3rd segment as in graminicola.
Legs black, apart from the tips of the front femora.
Ratio of the 2nd and 3rd costal segments (Fig. 35) 3—3.2, 2nd and 4th
3.2 to 4.5 (contrast graminicola): 4th segment relatively long. Veins pale: mem-
brane pale. Squama white with a white fringe. Wing-length g 2.1 mm, 2
2.6—2.8 mm.
Otherwise as for graminicola.
Male genitalia. Distance from the end of the aedeagal apodeme to the tip of
the aedeagal hood 0.75 mm. Aedeagal apodeme short and thick, strongly bent
along its length: its posterior end (after the attachment of the inner processes
of the postgonites) broadened into a large head 4 times the width of the rest of
the apodeme: this head occupies 1/4, of the whole apodeme. 9th sternite (Fig.
13d) completely asymmetrical, its left side about twice as long as its right: it
bears a large broad hypandrial apodeme about equal in length to the right (small)
side of the 9th sternite: the tip of the hypandrial apodeme reaches nearly 1/5 of
the length of the aedeagal apodeme. Pregonites large: the left one rather angulate:
the right folding over and joining with the phallophore. Postgonites conspicuous,
aot adpressed against the pregonites as in other species: inner process on left side
short and strong: on the right side the phallophore, inner processes of the post-
gonite and the pregonite are obscured and partially fused. Aedeagal hood broad,
deep in profile (as in graminicola).
Phallus (Fig. 13c) dominated by the enormous distiphallus which reaches back
beyond the base of the basiphallus. Basiphallus V-shaped, short and strong, the
left side broader than the right. Large anvil-shaped lobe on the left side at the
junction of the basiphallus and mesophallus: this hangs downwards and is larger
than in graminicola. Mesophallus developed into two large club-shaped appendages,
the left one larger, bilobed at its tip. A conspicuous sclerotised band joins the two
sides of the mesophallus across the top of the distiphallus. Distiphallus (Fig. 13b)
as in graminicola but larger, over 11/5 times the length of the basiphallus.
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 153
Ejaculatory bulb (Fig. 13a) fairly broad, rounded, with a clearly defined central
channel.
On external characters close to graminicola. The genitalia are however consider-
ably modified.
Larva. There are no larval specimens. However the following characters
have been described from the puparium with the type. Mandibles (Fig. 45a)
and paraclypeal phragma (Fig. 45b) as described for graminicola. Hind spiracles
(Fig. 46d) separated by 3—31/, times their own width, with elongate-oval bulbs:
adjacent bulbs forming an angle of 80° (contrast gramznicola). (More material
is needed to confirm the constancy of this last character.)
Biology. Mine (Fig. 53) on upperside of leaf. Eggs laid singly near the
leaf-margin, often several at different places on the same leaf. The resulting
larvae eat first towards the leaf-tip, then turn and head for the base of the leaf.
The mine gradually increases in width, but does not exceed 4 mm wide, remaining
semi-linear. Frass in large conspicuous particles.
Pupation follows outside the mine, the puparia adhering to the leaf near the
mine (conttast graminicola). Puparia reddish brown.
At least two generations. Larvae found in April produced flies between July
and September. This suggests that in the wild the puparia aestivate (as with many
Mediterranean species) and that there will be 2—3 generations during autumn,
winter and spring.
Material examined. In K. A. SPENCERS collection. Type & (with
preparation), Algeciras, Spain, 25.IV.55, ex Phragmites communis: emerged
6.VII.55. Allotype 9, same data, emerged 24.IX.55. Paratypes 5 9, same data,
emerged July-Sept. 55.
In the author’s collection. Paratype 9, same data, emenrgceds 22.1 epe
HERING (1951a) also observed this species at a neighbouring locality (as
graminicola).
Distribution. Southern Spain.
Host plant. Phragmites communis Trin.
Agromyza oryzae (Munakata, 1910)
Oscinis oryzae Munakata, 1910, Konchu Sekai, vol. 14, p. 7—11.
Oscinis oryzella Matsumura, 1915. Man. Inj. Ins. Japan, vol. 2, p. 53.
Agromyza oryzae Hendel, 1931, Flieg. pal. Reg., vol. 59, p. 142; etiam Kuwayama et Kato.
Agromyza oryzella Yuasa & Koyama, 1938, Bot. & Zool., vol. 6, p. 820.
Wexthicuresi15a5 15€, 26, 36
A fuller synonymy is given by Karo (1956) from which the above is abridged.
The following treatment owes much to that paper, which analysed the external
characters of 165 Japanese specimens.
Frons narrow, as broad as, or a little exceeding the eye-width at the level of the
front ocellus, parallel-sided. Antennae clearly separated at their bases. In profile
(Fig. 26) the cheeks and orbits clearly visible, forming a continuous ring around
154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 155
the eye. Eyes with scattered white hairs. Eye-height about 5 times the height of
the jowls. 2 strong upper orbital bristles directed backwards: 3—6 lower orbitals
directed inwards and standing closer together than the upper orbitals. Ocellar
bristles very small. 3rd antennal segment broadened, somewhat angular apically.
Head usually entirely black; sometimes frons and first two antennal segments
brownish.
Dorsocentrals strongly developed, normally 4 + 2 clearly visible in profile
(two being presutural). Acrostichals arranged in 6 irregular rows. Legs black or
dark-brown, sometimes the tips of the front femora and the front tarsi are
yellowish. Front tibiae and tarsi with outstanding hairs on their anterior edge.
Ratio of 2nd and 3rd costal segments (Fig. 36) about 4; 2nd and 4th about 5.
Wing-tip between the ends of m, +. and r44 5. Veins brown: membrane more
or less clear. Squama greyish or ochreous, with a black or dark-brown fringe.
Wing-length about 2.3 mm.
3 genitalia. Distance from the end of the aedeagal apodeme to the tip of the
aedeagal hood 0.8 mm. Aedeagal apodeme more or less evenly curved downwards,
heavily sclerotised. The strengthened part posterior to the attachment of the inner
processes of the postgonites occupies 1/, of the whole apodeme. 9th sternite
rather straight-sided, bearing a short indistinct hypandrial apodeme. The pre-
gonites are as figured for nigripes (Fig. 2): postgonites larger than those figured
for nigripes (Fig. 2). Aedeagal hood large, weakly sclerotised around its edges.
Phallophore not clearly indented on its posterior edge in lateral view (compare
albipennis, Fig.’ 1).
The basiphallus (Fig. 15c) consists of two rods, the left being obviously broader
than the right. Mesophallus without any clearly defined “thorn” (contrast
distorta): the right side forms a weakly sclerotised lobe. Distiphallus very short,
clearly shorter than the basiphallus.
Ejaculatory bulb (Fig. 15a) moderately sclerotised, except along its distal edge,
a little narrower than long: central channel visible, reaching almost to the distal
edge. Stem fairly short, weakly sclerotised.
Larva. The larva may be compared with that of distorta as follows. For a
more detailed description and figures consult KATO (1956).
Mandibles with 5 (rarely 4) alternating teeth (as figured for distorta). Para-
clypeal phragma largely pale brown. Patches of bristle-like spines on the locomotory
humps (Kriechwiilsten) of the mesothorax, metathorax and all abdominal segments.
Front spiracles with 88—133 bulbs, distributed over the entire surface of the
spiracle. Posterior spiracles much as figured for distorta, narrowly separated.
Biology. Mine on upperside of leaf. Eggs laid singly near the leaf-tip; the
resulting larva produces a gradually broadening mine, heading towards the base
Figs. 39—42. Larval parts of Agromyza: 39, albipennis Meigen; 40, lucida Hendel; 41,
hendeli sp. nov.; 42, phragmitidis Hendel. a — mandibles; b — larval mouthparts;
c — front spiracles; d — hind spiracles
156 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
of the leaf. Frass particles fairly conspicuous, scattered. Zig-zag feeding lines
visible. (Only one mine specimen was available, so I do not know if this is
typical).
Pupation follows outside the mine. There is puparial dimorphism as in distorta,
but not so strongly marked as in that species. KUWAYAMA (1950) noted the fol-
lowing important differences between the two types of puparia.
(In mm) Hibernant puparium Non-hibernant puparium
Mean length 2.598 2.855
Mean width 1.645 1.595
Mean height 1.542 1.385
Thickness of puparial skin 0.0294 0.0205
Colour Dull black, rarely Pale greenish brown, brown or
brownish black dark-brown, rarely blackish,
glossy.
Kato (1956) adds the following: ”Hibernant puparium: Generally all the
segments not polished as having many microscopic furrows or lumpy surface;
second to fifth abdominal segments more or less subshiny, each having a distinct
transverse keel girdling median position; sixth and seventh abdominal segments,
each with weakly developed keels on both lateral sides, having several longitudinal
and oblique keel-like wrinkles on dorsum: eighth abdominal segment generally
without median keel or distinct wrinkles: anterior spiracles protruded rather
laterad and the distal surface of each spiracle directed strongly or sometimes
almost entirely laterad. -
Non-hibernant puparium: Generally all the segments smooth and shiny, without
any distinct keel or wrinkle; last three segments sometimes more or less subshiny
according to the existence of microscopic furrows; anterior spiracles protruded
rather ventrad and the distal surface of each spiracle directed strongly or some-
times almost entirely ventrad.”
These differences correspond with those observed in the European distorta.
In that species the differences are even more marked. KUWAYAMA also observes
that the hibernant puparium ’’does not adhere closely to the rice-leaf”, while the
non-hibernant type does. This too is confirmed by my observation of distorta.
There are at least two generations. In Japan the first generation of larvae feed
in June. Further details of the life-history are not known to me, though they
are probably given in some of the Japanese papers listed by KATO (1956) in his
synonymy. The flies were taken freely in June.
Material examined. In the HENDEL collection. & (with preparation),
type of the synonymous Agr. oryzae Hendel, ex Oryza sativa, Nikolsk-Ussurisk,
Süd-Ussurigebiet, Siberia, 27.VII.28. |
Distribution. Northern Japan (Hokkaido, Tohoku and Hokuriku pro-
vinces), East Siberia (South-Ussuri province).
Host plants. Gramineae, tribe Oryzeae: Oryza sativa L., Zizania latifolia
Turcz.
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 157
43a 43d 44a
46d
Figs. 43—46. Larval parts of Agromyza: 43, spenceri sp. nov.; 44, prespana Spencer;
45, graminicola Hendel; 46, baetica sp. nov. a — mandibles; b — larval mouthparts;
d — hind spiracles
158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
Agromyza distorta Griffiths 1955
Agromyza distorta Griffiths, 1955, Ent. Gaz., vol. 6, p. 10—12.
Textfigures 14a, 14b, 14c, 25, 37, 47a, 47c, 47d, 47e, 47f
Frons very broad, 1.5—2 times the width of an eye at the level of the front
ocellus, parallel-sided. Antennae clearly separated at their bases. In profile
(Fig. 25) the cheeks and orbits clearly visible, forming a continuous ring
around the eye. Eyes with scattered white hairs. Ratio of the eye-height to the
height of the jowls 5—7; 4—5 strong orbital bristles, clearly differentiated into
upper and lower orbitals: the upper directed backwards, the lower inwards.
Ocellar bristles normal-sized, reaching beyond the bases of the 2nd pair of upper
orbitals. 3rd antennal segment broadened apically, rather short, somewhat angulate
at its apex. Colour of head entirely black, or at most the lunule and first antennal
segment brown.
Dorsocentrals strongly developed (as oryzae), 4 + 1 or 4 + 2 clearly visible
in lateral view, one or two presuturals being clearly developed. Acrostichals
8-rowed. Legs black, except for the yellow tips of the front femora. Front tibia
with strong hairs on its anterior edges (though not so marked as in oryzae).
Ratio of the 2nd to 3rd costal segments 2.7—3; 2nd to 4th 4—5. Wing-tip
between the ends of ry, „ and m, 4 9. Veins brown: membrane strongly brownish
tinged. Squama ochreous or dark grey with a dull black fringe. Wing length
2.5 mm.
g genitalia. Distance from the end of the aedeagal apodeme to the tip of the
aedeagal hood 0.9 mm. Aedeagal apodeme heavily sclerotised, somewhat bent
downwards; the strengthened part posterior to the attachment of the inner proces-
ses of the postgonites occupying a little less than 1/4 of the whole. 9th sternite
short and rounded (compare nigripes, Fig. 2), heavily sclerotised. Hypandrial
apodeme short, its tip reaching 1/5 of the length of the aedeagal apodeme.
Pregonites large (compare nigripes, Fig. 2): postgonites larger than those figured
for nigripes (Fig. 2), rounded. Aedeagal hood as for nigripes, but more weakly
sclerotised around its edges. Phallophore as figured for albipennis (Fig. 1).
Basiphallus (Fig. 14c) consisting of two rods of moderate width: mesophallus
asymmetrical: its left side developed into a black “thorn”, while its right forms
an ill-defined lobe of tissue. Distiphallus (Fig. 14b) very short, clearly shorter
than the basiphallus.
Ejaculatory bulb (Fig. 14a) drop-shaped, weakly sclerotised, almost transparent
in its distal 1/3: central channel clearly visible, reaching to 2/3 of the length of
the bulb: distal edge delicately rounded. Stem fairly short.
Larva. The following description is partly based on HERING (1956) who
gives a very detailed account of the larva.
Antennae and maxillary palpus small: longitudinal sclerite and mandibular
abductor apodeme absent. Mandibles (Fig. 47a) long, with 5 alternating teeth:
the upper two teeth longer than the lower three: right mandible higher than the
left. Distinct mandibular adductor apodeme present. No filaments above the
mandibles.
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 159
Labial sclerite slender: paraclypeal phragma strongly sclerotised.
Head bare. Prothoracic girdle consisting of about 8 rows of minute spines.
Metathorax bare. Dorsal edge of abdomen bare: strong bands laterally on the
metathorax and first three abdominal segments consisting of 14—18 rows of
spines: bands of the following segments decreasing progressively. Metathorax and
first three abdominal segments ventrally with a thick patch of elongate, bristle-
like spines on their locomotory humps (Kriechwülsten). 8th segment laterally,
7th and 8th ventrally, and the anal region with small fine spines.
Front spiracles (Fig. 47c) large and circular, bearing 200—250 bulbs distributed
over their whole surface: atrium short. Hind spiracles (Fig. 47d) projecting
somewhat, narrowly separated from each other, with 3 small egg-shaped bulbs:
adjacent bulbs forming an angle of about 90°.
Length (full-grown) about 5 mm.
Biology. Mine on upperside of leaf. Eggs laid singly near the edge of the
leaf. The larva first produces a long narrow channel towards the leaf-tip, then
turns and produces a large irregular blotch-mine. Frass particles scattered regularly
throughout the mine. Zig-zag feeding lines visible. Often several mines coalesce
to consume the whole leaf.
Pupation follows outside the mine. The winter puparia (Fig. 47e) are only
weakly attached to the leaf (the ventral surface not being flat) and easily fall into
the water.
The puparia are strongly dimorphic. The winter puparia (Fig. 47e) are black,
thick-shelled, strongly arched dorsally and obviously hollowed ventrally: the hind
end is very abruptly cut off. The summer puparium (Fig. 47f) is more elongate,
red, thinner-shelled, of fairly normal shape: for the group: dorsal edge only
moderately arched: ventral surface flat: hind end less abruptly cut off. The same
phenomenon occurs in oryzae, though to a lesser degree: detailed analyses by Ja-
panese workers are summarised under that species.
This species attacks large Glyceria plants growing in the water. The highly
modified front spiracles represent an adaptation to an aquatic environment (there
are several other instances in the Agromyzidae of startling changes in the structure
of the spiracles in species feeding on aquatic hosts). Nearly all the winter puparia
must end up in the water, as they are only very loosely attached to the leaf.
Furthermore I observed that one puparium (from which the type specimen later
emerged) was situated on a leaf which could only have been reached by the larva
crawling underwater. The patches of bristle-like spines on the ventral locomotory
humps clearly are a device to assist the larva in gripping a surface. These are
more strongly developed in oryzae.
Two generations. The larvae of the second generation are found in September.
Material examined. In the authors collection. Type g' (with pre-
paration) Ash Vale, Surrey, 5.IX.53, ex Glyceria maxima: emerged 18.V.54.
In the HERING collection (Berlin). & (with preparation), Aken, on the river
Elbe, with summer puparium, but no further data.
160 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
Distribution. England, Germany. Apparently a rare species. Not common
at the type locality.
Food plant. G/yceria maxima (Hartm.) Holmb.
Agromyza lunulata Sasakawa 1956
Agromyza lunulata Sasakawa, 1956, Sci. Rep. Saikyo Univ. Agr., vol. 8, p. 124—125.
This species was described from a single swept female. I have not attempted to
include it in this paper. SASAKAWA (1956) states that it is allied to distorta.
However until a male or the immature stages are found the question must remain
open. The external characters of the type certainly resemble distorta, but the third
antennal segment is clearly longer. Unfortunately no female of distorta is available
yet for comparison.
Fig. 47. Larval parts of Agromyza distorta Griffiths: a — mandibles; c — front spiracles;
d — rear spiracles; e — winter puparium; f — summer puparium (with front spiracles
missing)
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 161
The original description gives full details of the external morphology of the
type with figures of the head and wing.
PARASITES
The majority of Agromyzid parasites belong to the Chalcid family Eulophidae
and Braconids of the genus Opizs and the tribe Dacnusini. Species of each of
these groups occur as parasites of the #/gripes group. There is no evidence of
hyperparasitism.
One species of the Eulophidae has been bred freely from puparia of Agromyza
albipennis Mg. on Phalaris arundinacea. These Chalcids are unfortunately not yet
identifiable. The absence of Chalcid parasites from the majority of the species of
the n/gripes group is noteworthy. In the Poëmyza species, for instance, which also
feed on Gramineae the Chalcids form numerically the greater part of the parasites.
Several Optus spp. occur freely here. These too are not yet identifiable, but
this should be remedied soon, as Dr. MAX FISCHER of Vienna is working ou the
group, and has been sent much material bred from the Agromyzidae.
I can however give a reasonable account of the Dacnusini, which form the
majority of the parasites of this group. Some records were included in my paper
“Host Records of Dacnusini (Hym., Braconidae) from leaf-mining Diptera” in
1956. More can now be added. Mr. G. E. J. NIXON of the Commonwealth In-
stitute of Entomology has been kind enough to assist me in identifying these
species. All the specimens mentioned here have either been named by him, or by
myself by comparison with specimens named by him.
Antrusa melanocera (Thomson)
In GRIFFITHS (1956) this species was recorded as bred from albipennis, nigripes
and distorta. I can now add the following records.
Ex Agromyza nigripes Mg. on Glyceria fluitans and G. maxima collected at Ash Vale,
Surrey, 5 & 6.IX, 15 examples emerged in Sept./Oct. and April/May. Ex Agr. albipennis
Mg. on Phalaris arundinacea, same data and locality, 2 examples emerged 12.X and 14.III.
Ex Agr. distorta Grif. on Glyceria maxima, same data and locality, one example emerged
3.X. Ex Agr. lucida Hd. ex Glyceria maxima, same date and locality, one example emerged
19.III.61. Ex Agr. albipennis Mg. on Phalaris arundinacea, Woodwalton Fen, Hunts.,
22.VIII.60, one example emerged 4.IX.61.
These records add one further host, Agr. lucida Hd. The species is clearly
common on nigripes, but is less frequent on the related albipennis. At the Ash
Vale locality, where distorta and lucida are found feeding together with nigripes
on Glyceria maxima, Antrusa melanocera has occurred on all three species.
However only one example has been bred from /ucida, so that it seems probable
that gripes represents the main host, from which I have bred about 25 specimens.
The occurence of this species on distorta represents a clear case of disjunctive
cligophagy, for it does not occur on the Phragmites-feeding species which are more
closely related to nigripes than is distorta. This transference has clearly been
facilitated by the fact that distorta occurs on the same food-plant as nigripes.
162 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
Antrusa spec. a
Four examples of another Avtrusa species were bred from Agr. lucida Hd. on
Glyceria maxima, Ash Vale, Surrey, 5.IX.60, emerged 27.III—7.IV.61.
These specimens differ from typical melanocera in the more striate sculpture of
the petiole and the fewer hairs on tergite 3 + 4 and it is possible that they
represent a distinct species. One specimen of the normal melanocera was also bred
from the same host.
Dacnusa nydia Nixon
It has now become clear that this species, as formerly conceived, is composite.
The true »ydia is a parasite of nigripes though it is much less common than
Antrusa melanocera. Three examples were recorded by GRIFFITHS (1956).
Three more can now be recorded, bred from Agr. nigripes Mg. on Glyceria maxi-
ma, Ash Vale, Surrey, 5.IX.60, emerged 3.X.60 and March/April 1961.
Dacnusa sp. a (nydia group)
This species is represented by one specimen only bred from Agr. phragmitidis
Hd. on Phragmites communis, Slapton, S. Devon, emerged April °55 (K. A.
SPENCER).
Dacnusa spec. b (nydia group)
This species has been hitherto included under »ydia but can be easily recognised
by its possession of yellow tarsi with a contrasting black 5th segment, and its
broader petiole. My records are as follows:
Ex Agromyza hendeli spec. nov. on Phragmites communis, Woodwalton Fen, Hunts.,
23.VIII.60, 3 examples emerged 18—20.IX.60. Ex Agr. phragmitidis Hd. on Phragmites
communis, Woodwalton Fen, Hunts., 28.IX.60, 5 examples emerged April/May ’61. One
example from the same host and food-plant, Slapton, S. Devon, emerged April 55 (K. A.
SPENCER).
The species is clearly a common parasite of both the British Phragmites-feeding
species, hendeli and phragmitidis.
Dacnusa cincta Haliday
casions, the details being as follows:
Ex Agromyza lucida Hd. on Deschampsia caespitosa, Brookman’s Park, Herts., 6.VIII.54,
4 examples emerged 1.IX.54 and 9—10.V.55. Ex Agr. lucida Hd. on Glyceria maxima,
Ash Vale, Surrey, 5.IX.60, one example emerged 3.V.61.
Dacnusa pione Nixon
This is another large species related to cincta, of which only two Swedish |
examples have been hitherto known. I have now bred two specimens from Agr. |
phragmitidis Hd. on Phragmites communis, Woodwalton Fen, Hunts., Be
This large characteristic species has been bred from Agr. lucida on two oc
|
& 28.IX.60, emerged 10.V11.61.
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 163
The late emergence dates are noteworthy and suggest that the species may be
single brooded. In captivity the vast majority of Dacnusini emerge before mid-
May from overwintering puparia.
50
Figs. 48—50. Leaf-mines of Agromyza: 48, albipennis Meigen on Phalaris arundinacea:
49, nigripes Meigen on Glyceria maxima; 50, lucida Hendel on Glyceria maxima (natural
size)
Dacnusa lugubris Nixon
NIXON (1945) records several examples of this species as having been bred
by Hamm and BLAIR from “Agromyza nigripes on reeds”. Judging from the late
164 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
dates (September and October) on which the puparia were collected, these records
probably refer to Agr. phragmitidis Hd., and I have listed this species as a parasite
of that fly.
The species recorded in GRIFFITHS (1956) as lugubris from Phytomyza psendo-
bellebori Hendel has been re-examined by NIXON, who now states that it is
definitely not the true /wgubris. The species related to lugubris are a difficult
group which is clearly in need of revision.
Dacnusa spec. c (lugubris group)
One example of a form near /ugubris was bred from Agr. nigripes Mg. on
Glyceria maxima, Ash Vale, Surrey, 5.IX.60, emerged 2.III.61. It is improbable
that this is conspecific with the species bred from “reeds, referred to above as
lugubris.
Dacnusa ninella Nixon
Nixon (1945) refers to this species three examples bred by Hamm from
“Agromyza nigripes on reeds” from puparia taken in October near Oxford. It
was bred together with lugubris. The host is probably Agr. phragmitidis Hd. and
I have listed »inella as a parasite of that species.
Dacnusa spec. d (lugubris group)
One example of an obviously distinct Dacnusa spec., probably referable to the
lugubris group, was bred from Agr. distorta Grif. on Glyceria maxima, Ash Vale,
Surrey, 5.IX.54, emerged 26.IV.55.
Dacnusa spec. e
One example of a new Dacnusa species was bred by SPENCER from Agr. baetica
spec. nov. on Phragmites communis, Algeciras, S. Spain, 25.IV.55, emerged
25.V.55. As with most Dacnusini from the mediterranean area it is undescribed.
The following is the list of the known Dacnusine parasites from each species
of the nigripes group.
Agromyza albipennis Meigen Antrusa melanocera (Thomson)
A. nigripes Meigen Antrusa melanocera (Thomson)
Dacnusa nydia Nixon
» spec. ¢ (lugubris group)
A. lucida Hendel Antrusa melanocera (Thomson)
oy Sees A
Dacnusa cincta Haliday
A. hendeli spec. nov. Dacnusa spec. b (nydia group)
A. phragmitidis Hendel Dacnusa spec. a (nydia group)
» spec. & (nydia group)
ex prone Nixon
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 165
(
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I
Figs. 51—53. Leaf-mines of Agromyza: 51, communal mine of phragmitidis Hendel on
Phragmites communis (compare also spenceri sp. nov. and hendeli sp. nov.); 52, graminicola
Hendel on Phragmites communis; 53, baetica sp. nov. on Phragmites communis (with
young mines). Natural size
166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
Dacnusa lugubris Nixon
, hinella Nixon
A. baetica spec. nov. Dacnusa spec. e
A. distorta Griffiths Antrusa melanocera (Thomson)
Dacnusa spec. d (lugubris group)
The species with the widest range of hosts is Antrusa melanocera (see the
discussion under that species). Of the other species Dacnusa spec. b occurs on both
hendelt and phragmitidis, the two Phragmites-feeding species, while the other
species of Dacnusini are recorded from one host only. It is probable however that
some of the species bred from phragmitidis will also occur on hendeli.
Agromyza albipennis Mg. appears to have few parasites. The only Dacnusine
which has been bred from it is Antrusa melanocera, and that is far commoner on
nigripes. The flies can always be bred with the greatest of ease. The absence of
parasite records from other species is of course due to the absence of material.
ACKNOWLEDGEMENTS
This paper could not have been written without the cooperation of several in-
dividuals and Museums in sending me material. I wish to express my thanks to
the following: Professor Dr. E. M. HERING of the Humboldt University Museum
in Berlin, Dr. M. Brier of the Vienna Natural History Museum for the loan of
HENDEL's material, Overlärare Nits Rypen of Hälsingborg, Sweden, Mr. K. A.
SPENCER of Hampstead, London, Dr. J. T. Nowakowski of the Polish Academy
of Sciences, Professor SASAKAWA of Kyoto Prefectural University, Japan, Mr. F. D.
GOODLIFFE of Long Sutton, Hants., Dr. H. ANDERSSON of Lund University for the
loan of a ZETTERSTEDT type, Dr. K. W. HARDE of the State Natural History
Museum in Stuttgart for GROSCHKE's material and Mr. R. L. Cor of the British
Museum (Natural History) for the loan of a prespana paratype. My sister, Mrs.
E. A. OSBORNE, drafted some of the drawings.
SUMMARY
1. The characters of the nigripes group of the genus Agromyza Fallen are
defined. All the eleven Palaearctic species, including three, new to science, are
described and also two new subspecies.
2. The male genitalia of all species and subspecies are described and figured
for the first time.
3. The known larvae are described and figured: where larvae were not available,
larval characters have been reconstructed as far as possible from the puparia. An
attempt is made to rectify the considerable confusion which has reigned hitherto
over the identification of the larvae, and in particular the true »igrzpes larva is
described for the first time.
4. Details of the biology of all species are given as far as they are known.
5. The name /ucida Hendel 1920 is removed from the well-known Phragmites
species, to which it was wrongly applied by HENDEL himself, and returned to the
original species. The Phragmites-feeder is redescribed as hendeli spec. nov.
G. C. D. GRIFFITHS : Revision of Agromyza nigripes group 167
6. Detailed records are given of the distribution of all species. Two species,
albipennis and phragmitidis, extend right across the Palaeartic region to Japan:
three other species, nigripes, hendeli and lucida, are wide-spread in temperate
Europe. The known distribution of the other species is more restricted.
7. An attempt is made to clarify the host-relationship of albspennis, nigripes
and /ucida, which are recorded from many hosts.
8. Keys for the identification of adults and larvae are given.
9. Chapters on phylogenesis, host-relationship and previous workers’ treatment
of the group are included.
10. An account of the species of Dacnusini (Hym. Braconidae), parasitic on
this group, is given.
REFERENCES
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HENDEL, F., 1922. ,,Blattminierende Fliegen’. Wien. ent. Ztg., vol. 39, p. 65—72.
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HERING, M., 1927. ,,Minenstudien VIII”. Z. angew. Ent., vol. 13, p. 191.
HERING, E. M., 1951a. ‚Schwedische Miniaturen’. Opusc. ent. vol. 16, p. 81—88.
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HERING, E. M., 1955. „Agromyza nigrociliata Hendel als Getreideschädling (Dipt.)”.
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HERING, E. M., 1956. „Die Larven der Agromyziden (Diptera), II”. Tijdschr. Ent., vol. 98,
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HERING, E. M. 1957. „Bestimmungstabellen der Biattminen von Europa”. ’s-Gravenhage
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KALTENBACH, J. H., 1856. ,,Die deutschen Phytophagen aus der Klasse der Insekten, oder
Versuch einer Zusammenstellung des auf Deutschlands Pflanzen beobachteten
Bewohner und deren Feinde”. Verh. naturh. Ver. preuss. Rheinl. & Westph.,
vol. 13, p. 165—265.
KARL, O., 1926. ,,Minenzuchtergebnisse, I’, Stettin. ent. Ztg., vol. 87, p. 136—158.
Kato, S., 1956. "A taxonomic Study of the Rice Leaf-miner, Agromyza oryzae (Munakata)
(Dipt., Agromyzidae)”. Bull. nat. Inst. agric. Sci. (Japan), ser. C, vol. 6, p. 25—34.
KUWAYAMA, S., 1950. Proc. 8th Int. Cong. Ent. (Stockholm), p. 662.
MATSUMURA, S., 1915. "Dainippon Gaichu Zensho” (Manual of the injurious Insects of
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MEIGEN, J. W., 1930. „Systematische Beschreibung der bekannten Europäischen zweiflüge-
ligen Insekten’, vol. 6, p. 166—196.
168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 2, 1963
MEIJERE, J. H. C. DE, 1925. „Die Larven der Agromyziden’’. Tijdschr. Ent, vol. 68,
p. 195— 293.
Meijer, J. H. C. DE, 1928. Ibid, „Erster Nachtrag”. Idem, vol. 71, p. 145—178.
MEIJERE, J. H. C. DE, 1934. Ibid, „Zweiter Nachtrag”. Idem, vol. 77, p. 244—290.
MEIJERE, J. H. C. DE, 1940. Ibid., „Fünfter Nachtrag”. Idem, vol. 83, p. 161—188.
MEIJERE, J. C. H. DE, 1943. Ibid., „Siebenter Nachtrag”. Idem, vol. 86, p. 61—76.
MUNUKATA, T., 1910. „Konchu Sekai” (Insect World), vol. 14, part 9, p. 7—11. (in
Japanese)
NIXON, G. E. J., 1945. "A Revision of the European Dacnusini (Hym., Braconidae,
Dacnusinae)’’. Ent. mon. Mag., vol. 81, p. 189—229.
NIXON, G. E. J., 1946. Idem. Ibid., vol. 82, p. 279—300.
PHILLIPS, W. J., 1914. ’’Corn-leaf blotch Miner”. J. agric. Res., vol. 11, p. 15—31.
SASAKAWA, M., "New Agromyzidae from Japan XII”. Sci. Rep. Saikyo Univ., Agr., vol. 8,
os 1222 131.
SASAKAWA, M., 1958. "The Female Terminalia of the Agromyzidae, with Description of a
new Genus (I)”. Sci. Rep. Saikyo Univ., Agr., vol. 10, p. 133—150.
Sécuy, E., 1934. ’Dipt. (Brachycéres) Muscidae acalyptratae”. Faune Fr. vol. 28, p.
536—626.
SPENCER, K. A., 1956. "Observations on the British Agromyzidae (Dipt.). — II. Records
of Agromyzidae (Dipt.) in Scotland”. Ent. Gaz., vol. 7, p. 51—54.
SPENCER, K. A., 1957. "Two new European Species of Agromyzidae (Dipt.)”. Ent. mon.
Mas y:0l.293.2P255 37.
SPENCER, K. A., 1959. ”A Synopsis of the Ethiopian Agromyzidae (Dipt.)”. Trans. R.
ent. Soc. Lond., vol. 111, p. 237—329.
SPENCER, K. A., 1960. "Records of further Ethiopian Agromyzidae (Diptera), mainly from
South Africa, including eighteen Species new to Science”. Trans. R. ent. Soc.
Lond., vol. 112, pt. 2, p. 15—36.
STARY, P., 1930. „Über minierenden Insekten Mährens und Schlesiens’’. Acta Soc. Sci. nat.
Morav., vol. 6, fasc. 6, p. 125—242. (in Czech)
STROBL, G., 1893. „Beiträge zur Dipterenfauna des österreichischen Littorale, XV,
Muscidae, B. Acalypterae”. Wien. ent. Ztg., vol. 12, p. 121—136.
Yuasa, H. and Koyama, T., 1938. Bot & Zool., vol. 6, pt. 4, p. 820.
ZETTERSTEDT, J. W., 1840. Insecta Lapponica. Sectio III: Ordo V, Diptera, p. 783—794.
ZETTERSTEDT, J. W., 1848. Diptera Scandinaviae, vol. 7, p. 2728—2844.
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COMPARATIVE MORPHOLOGY OF THE SALDIDAE AND
MESOVELIIDAE (HETEROPTERA)*
BY
APR GUPT
Department of Entomology, University of Idaho, Moscow, Idaho, U.S.A.
CONTENTS
Rr Ero chu Gd Orne teen 169
Mate als An dsMetho dsnyne RI e ocio 169
NIR Comparativemdesctiptions annen 170
INARIM orpholoeicalstconsiderationssandmeompatisons Ws nee 180
We ATOM CASE VO NOTTE aun, fe nil Re Mae enc hn eis 194
MES Me IONI 194
VALUE Te ALU ORACLE IR TT 195
I. INTRODUCTION
There has been little agreement among heteropterists on the probable position
of the Saldidae in the higher classification of the Heteroptera, and this family,
more than any other, has been variously moved from one group to another.
Recently, on the basis of his studies of the female genitalia of the Heteroptera,
SCUDDER (1959) suggested that the Saldidae might be closely related to the Meso-
veliidae. A comparative morphological study of these two families was, therefore,
thought to be necessary to throw some light on the probable position of the Sal-
didae, and its relationship with the Mesoveliidae. Both alate and apterous forms of
the two families have been studied and their morphology is compared and dis-
cussed.
In addition, two general aspects of the morphology of the Heteroptera have
been considered namely, the interpretation of the head sclerites and the variation
in the thoracic structure between apterous and macropterous forms of the two
families.
II. MATERIALS AND METHODS
Saldula palustris (Douglas) was collected from the beach at Point Grey, Van-
couver, B.C. Aepophilus bonnairei Sign. from Jersey, Channel Islands, was kindly
supplied by Dr. G. G. E. SCUDDER of the University of British Columbia. Meso-
velia mulsanti White from England and Holland were obtained from Mr. I.
LANSBURY (Hope Department of Entomology, Oxford) and Dr. R. H. CoBBEN
*) Based in part on a thesis submitted in partial fulfilment of the requirements for the
degree of Master of Science of the University of British Columbia, Vancouver, B.C.
169
170 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
(Wageningen, Netherlands), respectively. The alate specimens of Mesovelia vitts-
gera Puton were obtained from the Musée Royal de L'Afrique Centrale, Tervu-
ren, Belgium, and were originally collected in the Congo. Other specimens of the
families belonging to the four higher taxonomic groups — Pentatomomorpha,
Cimicomorpha, Amphibicorisae and Hydrocorisae — were also used for compa-
rative study.
The external anatomy was studied both from dried and alcohol-preserved
specimens. The material was boiled in 10% potassium hydroxide, passed through
glacial acetic acid, stained in acid fuchsin, and cleared in creosote. Observations
on the cephalic muscles of Saldula were made from specimens preserved in 70%
alcohol, passed through different grades of alcohol, cleared in xylene and examined
under polarized light.
All drawings were made by using a squared graticule eye piece, and are not
made to the same scale.
III. COMPARATIVE DESCRIPTIONS
Head*)
Saldula palustris (Douglas) (Figs. 1—3)
Head short, broad, vertex well developed, with a lateral invagination near eye
extending anteriorly to frons; coronal and ecdysial cleavage lines distinct in
nymph, absent in adult; frons short, frontoclypeal sulcus absent; clypeal region
relatively extensive, differentiated into postclypeus, anteclypeus and paraclypeus,
cephalic portion of postclypeus with two lateral unpigmented areas; bucculae well
developed; postgenal bridge short; mandibular plate absent, mandibular lever well
developed, triangular (in nymph and adult); postocciput in the form of thin ring
around occipital foramen, apparently differentiated into dorsal and lateral ele-
ments, lateral parts bearing paired condyles; labrum broad and flap-like, reaching
distal end of second labial segment, epipharyngeal process absent; labium four-
jointed, first segment thickest, third longest, being swollen proximally and
tapering distally; antennae four-segmented with small intersegmental sclerites, |
first segment much thicker than rest, the whole beset with hairs, third and
fourth segments also bearing stout bristles; eight to ten pairs of trichobothria
present, scattered over vertex, frons, and postclypeus (nymph with four pairs, one
pair on frons, two pairs on postclypeus and fourth pair on anteclypeus); com-
pound eyes conspicuous, two ocelli present; two unpigmented areas present laterad
of ocelli.
Aepophilus bonnairei Sign. (Fig. 4)
Similar to Saldula in essential parts, but differing in following features. Ce-
phalic portion of postclypeus without two lateral unpigmented areas, paraclypeal
region not as well defined as in Saldula, restricted to upper two thirds of ante-
clypeus; maxillary plate area well developed; bucculae not well developed; labrum
*) The terminology of parts follows that of SNODGRASS (1960).
A. P. Gupta: Morphology of Saldidae and Mesoveliidae 171
Figs. 1—3. Head of Saldula. (1) dorsal (frontal) view; (2) lateral view; (3) ventral view.
Fig. 4. Head of Aepophilus bonnairei, dorsal view. Figs. 5—7. Head of Mesovelia mulsanti:
(5) dorsal view; (6) lateral view; (7) ventral view
reaching distal end of first labial segment, and not second as in Saldula; com-
pound eyes not conspicuous, ocelli absent; postocciput not divided into dorsal and
lateral elements; postgenal bridge longer than in Sa/dula; four pairs of trichobo-
thria — one pair on frons, two pairs on postclypeus and one pair on anteclypeus.
Mesovelia mulsanti White (Figs. 5—7)
Head longer than in Saldula; vertex well developed, and overlapped by pro-
thorax; coronal and frontal ecdysial cleavage lines indistinct in nymph and adult;
frontoclypeal sulcus absent; clypeal region well developed, differentiated into post-
clypeus, anteclypeus and paraclypeus; maxillary plate area well developed, sepa-
rated from paraclypeal region by a short genal sulcus; lower limit of maxillary
plate area delimited by an indistinct line; bucculae poorly developed; mandibular
plate absent, mandibular lever well developed, roughly quadrangular; labrum flap-
like, with an epipharyngeal process extending almost to distal end of second labial
segment; postgenal bridge longer than in Saldula; postocciput indistinguishable
from occiput, and bearing two dorso-lateral condyles; labrum four-jointed; first
segment thickest, third longest, being swollen on inner side proximally and tapering
distally; antennae four-jointed with small intersegmental sclerites, first antennal
1972 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
segment thicker than rest, bearing a stout bristle lateromedially; three pairs of
trichobothria, one pair on frons, two pairs on postclypeus (in the nymph one ad-
ditional pair on anteclypeus); compound eyes inconspicous, ocelli rudimentary.
Mesovelia vittigera Puton
Similar to M. mulsanti but ocelli well developed.
Thorax
Saldula palustris (Douglas) (Figs. 8—13, 30—34)
Prothorax. Pronotum large, with anterior collar, posteriorly overlapping base of
fore wings, callal area broad, dome-shaped and triangularly depressed in middle;
episternum shorter than epimeron, forming a precoxal shelf, pleural sulcus and
pleurodema distinct but short; trochantin short, very distinct in nymph; sternum
differentiated into a transversely elongated presternum, a triangular basisternum,
and a sternellum, furcal arms elongate, extending laterally to meet pleurodema.
Mesothorax. Mesonotum differentiated into scutum and scutellum, with dividing |
sulcus incomplete, scutum secondarily divided into a median and two lateral areas,
scutellum extending posteriorly over second abdominal segment, and apically |
pointed, parascutellum extending anterolaterally into scutoscutellar region, the
latter being unequally sclerotized, postscutellum visible laterally; pleural sulcus |
distinct, short, pleurodema short, directed anteriorly; trochantin short; episternum
forming precoxal shelves, epimeron divided into a dorsal anepimeron and a ventral
katepimeron, the latter produced into a point at lower limit of coxal cleft; pleural
wing process very distinct; basalar and subalar sclerites not distinguishable; ster-
num well developed, and divided into a presternum, a basisternum, and a sternel-
lum, the latter extending posteriorly over metasternum; furca well developed, with
furcal arms extending laterally and meeting pleurodema; phragma well developed.
Metathorax. Metanotum in the form of fused metascutum and scutellum, post-
scutellum well defined; pleural sulcus horizontal, and dorsal in position, pleuro-
dema short; trochantin comparatively long; episternum broad, forming a large
precoxal shelf, two precoxal shelves being approximated medially, epimeron
small and dorsal in position; orifice of scent-apparatus located laterally; basalar
and subalar sclerites indistinguishable; metapleural wing process lying forward in
region of mesepimeron, and reinforced by processes from postscutellum and
epimeron of mesothorax and postscutellum of metathorax; metasternum reduced to
a small plate beneath mesoscutellum; furcal arms short, not reaching pleurodema;
phragma very large.
Wings (Figs. 30, 31)
Fore wings differentiated into clavus, corium, embolium, and a membranous
area, posterior end of clavus produced medially into a narrow triangular area along
inner margin of membrane, membranous area with four cells.
Hind wings with distinct vannal and jugal folds, costa, subcosta and base of
A. P. Gupta: Morphology of Saldidae and Mesoveliidae 173
2
Figs. 8—14. Thorax of Saldula: (8) dorsal view; (9) ventral view; (10) lateral view of
prothorax; (11) lateral view of pterothorax; (12) internal view of prothorax showing
pleurodema and apodemes; (13) internal view of mesothorax showing furca, phragmata and
pleurodema; (14) internal view of metathorax showing furca, phragmata and pleurodema
radius fused, radius and media distally fused, base of media and cubitus in
contact with distal median plate, two vannal veins in the vannal area, 2V with a
thickened base, jugal lobe with single jugal vein; humeral plate well developed,
first and second axillaries reduced, third axillary articulated both with proximal
median plate and second axillary, two median plates (proximal and distal)
present.
Legs (Figs. 32—34)
Coxae, especially those of hind leg, well developed, having only pleural and
trochantinal articulations, hind coxa with inflected articular surface, proximal half
174 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
with a coxal suture, internal ridge of suture continuous with inflection of articular
surface, fore and middle coxae without coxal suture, distally coxae bearing anterior
and posterior articular surfaces for articulation with trochanter, trochanter immo-
vably articulated with base of femur; femora flattened laterally; hind tibia longer
than hind femora, bearing stout bristles, proximal end with a distinct head bent
toward femur; tarsi consisting of three tarsomeres, the basal the smallest, tar-
someres with bristles, pretarsus in the form of two claws.
Aepophilus bonnairei Sign. (Figs. 15—17)
Prothorax. Pronotum large with a pronotal collar; epimeron broader than
episternum, precoxal shelf formed by both; pleural sulcus and pleurodema short;
coxal cleft prominent, trochantin present; sternum differentiated into a basister-
num and sternellum; furcal arms short.
Mesothorax. Mesonotum a triangular piece; pleural sulcus short (shorter than
that of prothorax); trochantin present; sternum differentiated into a presternum,
basisternum and sternellum, latter extending over metasternum (as in Saldula) ;
furca not well developed.
Metathorax. Metanotum undifferentiated, shorter than pronotum and meso-
notum; pleural sulcus dorsal in position; epimeron dorsal; pleurodema very short;
episternum broad, forming a large precoxal shelf.
Legs. Coxae more or less like those of Saldula, no outer suture and inflection,
anterior and posterior articular processes present, femora flattened laterally, hind
tibiae longer than hind femora as in Saldula, tibiae with stout bristles at their
distal ends, tarsi with three tarsomeres.
Two rudimentary mesothoracic wings present.
Mesovelia mulsanti White (Figs. 18—23)
Prothorax. Pronotum large, with a short collar; epimeron broader than epister-
num, precoxal shelf formed by both episternum and epimeron; pleural sulcus in-
distinguishable, a very short pleurodema present; trochantin present, very distinct
in nymph; sternum differentiated into a basisternum and a sternellum; furcal arms
(apophyseal apodemes) small, pointing ventro-posteriorly.
Mesothorax. Mesonotum undifferentiated; pleural sulcus indistinguishable, very
short pleurodema present; episternum medially fused with basisternum; trochantin
present; precoxal shelf formed by both episternum and epimeron; sternum diffe-
rentiated into basisternum and sternellum; furcal arms (apophyseal apodemes)
short, pointing posteriorly.
Metathorax. Metanotum undifferentiated; pleural sulcus indistinguishable; coxal
cleft absent (present in prothorax and mesothorax); apophyseal apodemes short,
pointing laterally; episternum fused with basisternum, forming a part of precoxal
shelf; metasternum differentiated into basisternum and sternellum, latter fused
with first abdominal sternite; single median orifice of scent apparatus present.
Legs. Coxae well developed, fore and middle coxae being swollen, articulation
both pleural and trochantinal, hind coxae without coxal sutures on their outer
A. P. Gupta: Morphology of Saldidae and Mesoveliidae 175
PNC
Figs. 15—17. Thorax of Aepophilus bonnairei: (15) dorsal view; (16) ventral view; (17)
lateral view
surfaces; trochanter immovably attached to base of femur, femora flattened
laterally, fore and middle femora with more stout bristles than hind femora, hind
tibiae with stout bristles, middle tibiae with combs at their distal ends, three
tarsomeres, pretarsus in the form of claws, latter provided with pseudoaroliae
(parempodium).
Mesovelia vittigera Puton (Figs. 24—29)
Prothorax. Pronotum with anterior collar, posteriorly overlapping bases of fore
and hind wings, pronotum with a wide callal area, the latter with two laterally
176 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
located depressions; epimeron broader than episternum; pleural sulcus short,
pleurodema very short; trochantin present; sternum as in M. mulsanti.
Mesothorax. Mesonotum differentiated into scutum and scutellum, the latter
not extending posteriorly over second abdominal segment as in Saldula; post-
scutellum present; sternopleural region the same as in M. mulsanti.
Metathorax. Metascutum and scutellum fused, median part extending over
second abdominal segment; postscutellum distinguishable; pleurosternal region the
same as in M. mulsantz.
Wings (Figs. 28, 29)
Fore wings with clavus, corium, an indistinct embolium, bases of costa, sub-
costa, radius fused, distal ends of fused costa, subcosta and radius, and media and
cubitus form a stigma, rest of wing membranous, but without cells.
Hind wings without jugal fold, vannal fold present, bases of costa, subcosta,
radius and media fused, radius and media distally fused, two vannal veins present.
Legs. As in M. mulsantz.
Abdomen
Saldula palustris (Douglas) (Figs. 35—39, 45, 46, 50)
Abdomen with ten segments. Terga of segments two to eight differentiated
into dorsal median plates and lateral paratergites, tergum of first segment not
complete medially, in the male posterior margin of second paratergite modified
into a granulated convex lobe; conjunctiva between second and third paratergite
forming beneath the anterior margin of third paratergite a fold with a concave
anterior margin, the latter being beset with stout setae; this together with convex
lobe of second paratergite forms, during copulation, grasping mechanism in male;
remnants of orifices of abdominal scent-glands present at posterior margin of
third tergite.
Sternum of first segment rudimentary and indistinguishably fused with partially
membranous sternum of second segment; seven pairs of spiracles present on
lateral margins of sterna two to eight.
Female genitalia (Figs. 36—39)
Previous descriptions: EKBLOM (1926); LESTON (1956); SCUDDER (1959).
First gonapophyses joined by membrane, elongate, tapering and split longitu-
dinally with serrate dorsal tips, rami sclerotized; first gonocoxa fused with ventral
part of eighth paratergite; ninth tergum without separate paratergite; gonangulum
triangular, its anterior limbs being fused with first ramus, and posterior side with
ninth tergum, ventral angle of posterior side articulating in a notch on middorsal
side of second gonocoxa, thus forming a fulcrum on which the latter pivots;
second gonapophyses broader than first, sclerotized and united except at apices,
the latter being truncate; second gonocoxa elongate and thickened dorsally; gono-
A. P. Guerra: Morphology of Saldidae and Mesoveliidae 10797
FM
TB
22 23
2
Figs. 18—20. Thorax of Mesovelia mulsanti: (18) dorsal view; (19) ventral view; (20)
lateral view. Figs. 21—23. Legs of Mesovelia mulsanti: (21) fore leg; (22) middle leg;
(23) hind leg
placs broad, curved and free distally, united proximally by partially sclerotized
membrane; spermatheca single, with an oval bulb or receptacle, and an elongated
spermathecal tube; the latter communicating with vagina, a muscular pump with
a single flange present between receptacle and main part of duct; wall of vagina
lined internally with wrinkled chitinous intima, and strenghtened by a sclerotized
ring.
Male genitalia (Figs. 45, 46, 50)
Previous descriptions: EKBLOM (1926); MARKS (1951); PRUTHI (1925).
Aedeagus differentiated into phallosoma and endosoma, the latter being further
178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
divisible into conjunctiva and vesica; proximal part of phallosoma wide and mem-
branous, distal part being narrow, heavily sclerotized and bent over proximal part,
mouth of phallosoma located ventro-laterally and confined to its distal half, two
minute posteriorly-directed appendages located one on each side of the anterior
end of mouth, base of phallosoma supported by an inverted Y-shaped sclerite,
with its upper limb fused with junction of basal plates; dorsal half of conjunctiva
sclerotized, forming a curved sclerite with its two ends produced into lateral
“wings” on either side of mid-dorsal line, distal end of conjunctiva containing two
pairs of dorsolaterally located appendages, anterior pair smaller than posterior one,
ventral wall of conjunctiva containing near mouth of phallosoma a V-shaped
structure with two pairs of processes projecting anteriorly from inner angle of
V, inner pair being smaller than outer; vesica narrow, coiled, and in close contact
with ejaculatory duct; ejaculatory reservoir located at junction of conjunctiva and
vesica; basal plates completely fused in middle line, forming horseshoe-shaped
structure; capitate processes attached on to ends of basal plates; parameres long,
sickle-shaped, and pointed at apices, proximal ends being broader and curved for
muscle attachment.
Aepophilus bonnairei Sign. (Fig. 47)
Abdomen with ten segments. Terga two to eight differentiated into median
tergites and lateral paratergites; first tergum fused with second; clasping organ
present in second and third segments.
First sternum rudimentary and indistinguishably fused with partially mem-
branous second sternum; seven pairs of spiracles on segments two to eight.
Female genitalia. More or less as in Saldula palustris.
Male genitalia (Fig. 47). Aedeagus differentiated into a phallosoma and en-
dosoma, the latter with two appendages projecting out of mouth of phallosoma,
base of phallosoma membranous, distal part narrow and heavily sclerotized, bent
over proximal part, base of phallosoma supported by an L-shaped sclerotized
structure, horizontal limb being fused with junction of basal plates; dorsal part of
endosoma presents a sclerotized structure; basal plates completely fused and like
those in Saldula.
Mesovelia mulsanti White (Figs. 40—44, 48, 49, 51)
Abdomen with ten segments. Terga of segments two to eight in female and
two to seven in male differentiated into dorsal median tergites and lateral para-
tergites; orifice of abdominal scent-gland in middle of fourth tergum; clasping
organ in male absent.
First sternum indistinguishably fused anteriorly with metasternellum and post-
eriorly with second sternum; seven pairs of spiracles present on segments two to
eight; anterior margin of seventh sternum produced anterolaterally in mid-line
as a long apodeme; two circular patches of black setae present on eighth sternum
in male.
A. P. Gupta: Morphology of Saldidae and Mesoveliidae 179
Figs. 24—27. Thorax of Mesovelia vittigera: (24) dorsal view; (25) ventral view; (26)
lateral view of prothorax; (27) lateral view of pterothorax. Figs. 28—29. Wings of Meso-
velia vittigera: (28) fore wing; (29) hind wing
Female genitalia (Figs. 41—44)
Previous description: EKBLOM (1926); NEERING (1954); PENDERGRAST
(1957); SCUDDER (1959).
First gonapophyses joined by membrane, elongate, tapering and split longitu-
dinally, with serrate tips; rami sclerotized and interlocking; first gonocoxa fused
with eighth paratergite; gonangulum triangular, its posterior side also fused with
an inflection between eighth and ninth terga; second gonapophyses elongate, scle-
180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
rotized, laciniate, united except at tips; second gonocoxa elongate; gonoplacs trian-
gular, sclerotized, curved and attached to second gonocoxae; spermatheca single
with an accessory fecundation canal.
Male genitalia (Figs. 48, 49, 51)
Previous description: EKBLOM (1926); PRUTHI (1925).
Aedeagus differentiated into phallosoma and endosoma, the latter being further
divisible into conjunctiva and vesica; proximal part of phallosoma membranous,
distal part heavily sclerotized and bent over proximal part; a triangular sclerotized
area present in mid-dorsal part of conjunctiva, proximally conjunctiva produced
into sclerotized appendages projecting from mouth of phallosoma; vesica narrow
and short, in close contact with ejaculatory duct; two ejaculatory reservoirs
distinguishable; basal plates completely fused in middle line, forming a horseshoe-
shaped structure; capitate processes attached laterally; parameres hook-like, pointed
at apices, proximal ends being broader for muscle attachment.
Mesovelia vittigera Puton
Abdomen very similar to that of M. mulsanti.
IV. MORPHOLOGICAL CONSIDERATIONS AND COMPARISONS
Head
Saldula palustris (Douglas). In the adult insect, on the anterodorsal part of the
head capsule, the position of the frontal ecdysial cleavage line can be recognized
by the position of the two pits; these are found on each side of the vertex along
the frontal ecdysial cleavage line in the nymph. These two lateral pits are not
homologous with the pretentoria of Cicadellidae (SPOONER, 1938). They have
no counterparts in other Heteroptera as far as is known, and the name epicranial
pits is here suggested for these structures. It should be mentioned that the frons
is not always delimited laterally by the ecdysial cleavage line, for SNODGRASS
(1960) states, “they vary greatly in their extent and position in different insects.”
He suggested the name cephalic apotome for the part cut out at ecdysis by the
ecdysial cleavage line.
In Saldula the fronto-clypeal sulcus is absent, and consequently there is no
external demarcation between the frons and the clypeus. The positions of the
antafossae or the mandibular levers are often used as landmarks to delimit the
frontal and clypeal areas (SPOONER, 1938). However, in Saldula they are situated
cephalad and so are perhaps unreliable. These areas can be distinguished, however,
by muscle attachment; the dilator muscles of the sucking pump are attached to the
clypeus internally (SNODGRASS, 1935). SNODGRASS (1960) stated that the ci-
barium has often been called the ‘pharynx’ although it lies outside the mouth. He
further stated that this cibarium has become the sucking pump of the liquid-
feeding insects. It is evident, therefore, that the term pharyngeal pump is in-
A. P. Gupta: Morphology of Saldidae and Mesoveliidae 181
CO EB
HP
TAR
Figs. 30—31. Wings of Saldula: (30) fore wing; (31) hind wing. Figs. 32—34. Legs of
Saldula: (32) fore leg; (33) middle leg; (34) hind leg.
accurate. In Saldula the criterion of the muscle attachment has been used to
delimit the frontal and the clypeal areas, although it is realized that this proce-
dure is open to some criticism. For example, FERRIS (1944), pu PORTE
(1946), and PARSONS (1959) have criticized the criterion of muscle attachment
for the identification of the sclerites. The latter in her account of Gelastocoris used
the frontal ganglion as landmark to delimit the point of attachment of the dilator
muscle of the food-pump. She recognized two sets of muscles: cibarial, which lie
182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
posterior to the frontal ganglion and attach on the clypeus, and pharyngeal
muscles, which lie posterior to the frontal ganglion and attach on the frons. In
this she followed MARKS (1959). PARSONS (1959), however, suggests, “that the
muscle posterior to the frontal ganglion could come to insert on a cibarial portion
of the food-pump, or that cibarial muscles might shift their insertions to the
pharyngeal portion.” She further quoted MARKS (1959) and stated that the
position of the frontal ganglion also varies relative to the muscles from one
species to another. It is evident then that an interpretation based on the position
of the frontal ganglion is neither reliable. In the present study, therefore, in the
absence of any alternative, the criterion of muscle attachment has been adopted
to delimit the frontal and the clypeal portions of the head capsule. Examination
shows that the dilator muscles of the sucking pump attach to a definite part of
the head capsule, and so it is probably correct to interpret this area as the clypeus.
EKBLOM (1926) states that in Saldula saltatoria (Lat.), “the forehead forms in
front a transverse ridge where it limits the clypeus.” This ridge would appear to
be merely the posterior margins of the unpigmented postclypeal areas. In Saldula,
therefore, as in other Heteroptera, the clypeus appears to be differentiated into
a postclypeus, which is united with the frons, a distinct anteclypeal region, and
two paraclypeal lobes.
The anteclypeus is a clearly defined area, and appears externally as a convex lobe,
its internal concavity together with its lateral invaginations form a supporting base
for the food-pump. To the cephalic margin of the anteclypeus is attached the
flap-like labrum.
The paraclypeal lobes are well defined areas on each side of the anteclypeus;
they extend caudad up to half the length of the latter. They are clearly visible in
both the nymph and the adult, and appear to have no internal muscle attachment.
The homology of the paraclypeal lobes in the Hemiptera is very controversial, and
has been the subject of much debate. SMITH (1892) and WEBER (1929) con-
sidered them to be homologous with the mandibular plates. SNODGRASS (1935)
also mentions, “that these paraclypeal lobes appear to be the mandibular plates of
the Homoptera,” but he himself doubted their mandibular origin in the Homo-
ptera in the absence of any embryological evidence. EKBLOM (1926) and COBBEN
(1960) designated these lobes in Saldula saltatoria as “laminae maxillares’’ and
“maxillare Platten’ respectively. PARSONS (1959) stated that the paraclypeus in
Gelastocoris is wholly inflected within the head. However, most authors consider
the paraclypeal lobes as parts of the clypeus. Muir & KERSHAW (1911, 1912)
regarded them as “extensions of the clypeus”, and confirmed that “they have no
relation to the mandibles.” SFOONER (1938) stated that the paraclypeal lobes
are undoubtedly parts of the clypeus. MacGir (1947) referred to these two lobes
in Dysdercus intermedius Distant as juga.
It is usually not possible to trace the mandibular plates in the Heteroptera since
there is no sulcus between the mandibular plate area and the gena. It is, therefore,
advisable to consider the whole area of the head capsule between the eye and the
points of attachment of the mouthparts as the genal area; if mandibular plate area
need to be recognized, it is suggested that it be defined as the ventro-anterior area
of the gena to which the mandible articulates.
A. P. Gupta: Morphology of Saldidae and Mesoveliidae 183
Figs. 35— 39. Abdomen and female genitalia of Saldula: (35) dorsal view of abdomen of
male; (36) ventral view of female terminalia; (37) first gonocoxa and associated parts;
(38) second gonocoxa and associated parts; (39) spermatheca
In mandibulate insects the maxillae are usually attached to the ventral part of
the postocciput (SNODGRASS, 1935). It does not appear to be necessary to recog-
nize a maxillary plate area despite the fact that such an area is usually described
in the hemipterous head. Maxillary plates are absent in the primitive orthopteran
type of the head, and usually cannot be defined by sulci in the hemipterous head.
The origin of what is herein termed the postgenal bridge has been a problem
in the past. Many authors claim that the ventral region of the head is formed by
184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
the fusion ventrally of the maxillary plates, the area considered to be equivalent
to the postero-ventral part of the gena anterior to the point of articulation of
the labium. MACGILL (1947) referred to the ventral area of the head in Dysdercus
intermedins as a “large median ventral sclerite interpolated between the labium
and the foramen magnum.” However, she made no comment regarding its origin.
PARSONS (1959), like most earlier authors, adopted the term “gula” but noted that
perhaps it is not a true gula. SNODGRASS (1960) has recently considered the
ventral sclerites of the head in insects, and has concluded that they are not homo-
logous in all groups, and thus cannot in all be termed a “gula”. He mentioned
three distinct processes which may result in the formation of ventral sclerites of
the head in insects: in the first, a hypostomal bridge may be formed between the
occipital foramen and the base of the labium by the ventral fusion of two hypo-
stomal lobes as in the Diptera. The hypostomal bridge, he stated, is continuous
dorsally with the postocciput. The second modification, according to SNODGRASS,
is the ventral fusion of two postgenal lobes forming a postgenal bridge between
the occipital foramen and the base of the labium, as in Vespula maculata. When
this is the case, the hypostomal bridge is replaced by the postgenal bridge, which
differs from the former in not being continuous dorsally with postocciput. He
thus regarded the ventral plate in Notonecta and Naucoris as the postgenal bridge
since it is continuous with the postgenae and not with the postocciput. The third
process described by SNODGRASS is the ventral fusion of the lower ends of the
postocciput to form a median plate, which may become extended distally as in
Coleoptera. This median plate is the true gula, and is continuous proximally with
the postocciput.
In Saldula as in Notonecta and Naucoris, the ventral plate seems to be formed
by the fusion ventrally of the postgenae. However, a developmental study of
this region is badly needed to determine whether or not the homology accepted
in the present study is really the correct one. This study should include a con-
sideration of the origin of the bucculae. These structures, which apparently serve
to support the rostrum during feeding, are interpreted as the ventral extensions of
the areas anterior to the point of attachment of the labium. Since they often extend
posterior to the rostrum, this interpretation 1s perhaps incorrect.
Thorax
The prothorax in Saldula is more or less of a generalized type seen in other
Heteroptera in that it shows no differentiation into separate sclerites. The presence
of the dome-shaped callal area does not seem to be a constant feature in the
Saldidae, for DRAKE « CHAPMAN (1958) stated that a callus is absent in the
genus Saldoida.
Of the three thoracic segments, the mesothorax in Saldula is the most developed.
This agrees with WEBER’s (1930) thesis that in the Hemiptera the fore wings
are the principal organs of flight. Both the mesoscutellum and mesosternum are
well developed. TAYLOR (1918) stated that in the Heteroptera the mesothora-
cic sternum is indistinguishably fused with the pleura. This is true in Saldula, and
it is because of this fusion that the limits of the pleural and sternal sclerites cannot
A. P. Gupta: Morphology of Saldidae and Mesoveliidae 185
SSB
do
e
O 6)
x Co
55
—
4
ED)
47 > 43
Figs. 40— 44. Abdomen and female genitalia of Mesovelia mulsanti: (40) dorsal view of
abdomen of female; (41) ventral view of female terminalia; (42) first gonocoxa and
associated parts; (43) second gonocoxa and associated parts; (44) spermatheca
be clearly defined. The fusion of the sternopleural sclerotizations is also seen in
the metathorax. BRINDLEY (1934) figured the thorax of Saldula pilosella
(Thomson), and labeled the anterior and posterior areas of the mesosternum BS2
(basisternum of mesothorax) and BS3 (basisternum of metathorax), and left the
middle area unnamed. If the location of the apophyseal pits is taken as the
landmark in delimiting the sternal plates, her interpretation appears to be inaccu-
rate with respect to the mesosternum. According to the interpretation given, the
areas which she called BS2 and BS3 are merely the mesothoracic, presternum and
186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
sternellum respectively. The basisternum of the metathorax in Saldula is a small
plate, and lies beneath the mesosternellum, with the metasternal apophyseal pits
lying on each side of it. This can be seen if the mesothorax and the metathorax
are pulled apart. This structure is most clear in the nymph. Thus what BRINDLEY
considered as the basisternum of the metathorax is evidently the sternellum of
the mesothorax.
The pleural sulcus of the metathorax is horizontal, and lies on the margin of
the pleura, so much so that the epimeron occupies a dorsal position, and is at-
tached to the lateral margin of the metascutellum. According to TAYLOR (1918),
the horizontal position of the pleural sulcus seems to be a general feature in the
Heteroptera. BRINDLEY (1934) stated that this horizontal position is characteristic
of the aquatic bugs because of the enlarged coxae, which extend behind rather
than from beneath the thorax. LARSEN (1945) mentioned a horizontal pleural
ridge in Salda muelleri (Gmelin). The upper forward margin of the metapleuron
extends forwards beneath the posterior margin of the mesopleuron, and terminates
in the metapleural wing process in the region of the mesepimeron. TAYLOR
(1918) mentioned that similar condition is seen in the Nabidae, Gerridae and
Berytidae. However, the metapleural wing process in Saldula is peculiar in that
it is also supported by processes from the mesepimeron and the metascutellum,
and lies in the region of the mesothorax. The large precoxal shelves of the meta-
pleuron seem to be associated with the greatest development of the hind coxae.
In the fore wing, the corium is secondarily divided into a marginal embolium.
DRAKE and CHAPMAN (1958) also mentioned an embolium in the genus Sal-
doida. In the interpretation of the veins of the hind wing of Saldula the criterion
of the axillaries and their association with particular veins has been adopted. Thus
the veins in association with the distal median plate have been designated as the
media and cubitus, the former together with the radius seem to be very much
approximated toward the costa and subcosta. It may be mentioned here that dif-
ferent degrees of fusion of the costa, subcosta, and radius occur also in the
Miridae, Lygaeidae, Phymatidae, Mesoveliidae, and Piesmidae (HOKE, 1926).
DRAKE and Davis (1958) have figured the hind wing of the piesmid Mespa
splendida Drake, and have shown the cubitus in the same position as in Saldula.
HOKkE (1926) has studied the venation of the hind wings of the Heteroptera
in detail. She followed the COMSTOCK-NEEDHAM system, and divided the wing
into four areas: the costal area with costa, subcosta, radius and media 1 and 2;
medial area with media 3 and 4; cubital area with cubitus and first anal, and the
anal area with the remaining anal veins. On comparison, it becomes obvious that
her cubital area in Salda bouchervillei Prov. (= coriacea Uhler) and Saldula pal-
lipes (F.) (= separata Uhler) should really be interpreted as the vannal area, and
thus it appears that the vein which she designated as cubitus is probably one of
the vannal veins (Table I). Although she has omitted the consideration of the
axillaries in her interpretation, she has figured them in the two saldids she
studied, and it is evident that her cubitus is not in association with the distal
median plate, which it should be according to the interpretation (SNODGRASS,
1935) adopted in the present study. Since many of the veins of the adult wing in
the Heteroptera are without a corresponding trachea (HOKE, 1926) the inter-
A. P. Gupta: Morphology of Saldidae and Mesoveltidae 187
IO) SI
Figs. 45—46 and 50. Male genitalia of Saldula: (45) lateral view of aedeagus and as-
sociated structures; (46) frontal view of aedeagus and associated structures; (50) paramere
Fig. 47. Aepophilus bonnairei: lateral view of aedeagus and associated parts. Figs. 48—49
and 51. Male genitalia of Mesovelia mulsanti: (48) lateral view of aedeagus and associated
parts; (49) frontal view of aedeagus and associated parts; (51) paramere
pretation of the adult veins seems to be more reliable if based on the associated
axillaries. Very little information is obtained by studying the position of the
trachea in the nymph. A reinterpretation of the wing venation of the Heteroptera
based on the association of veins with axillaries is thus needed.
188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
TABLE I. INTERPRETATION OF THE VEINS OF THE HIND WING
HOKE GUPTA
Costa Costa
Subcosta Subcosta
Radius Media
rm m-cu
Media 1 plus 2 Cubitus
Cubitus Vannal 1
Anal 1 Vannal 2
Anal 2 Jugal 1
Abdomen
DRAKE and HOTTES (1951) stated that "the hardened and plate-like and
roughened lobe of the first paratergite together with the “peg-like” or “spine-
like” organs of the second paratergite form the stridulatory organ in the Sal-
didae. LESTON (1957), however, stated that the granulated plate and the pegs
were actually situated on the second and third paratergites respectively, and not
on the first and second as indicated by DRAKE and HOTTES. LESTON also stated
that the organ functions not as a stridulator but as clasping mechanism in the
male during copulation. Examination of the organ in Saldula has revealed,
however, that one small modification is needed in LESTON’s description; the
pegs are located actually on a fold of the conjunctiva underneath the anterior
margin of the third paratergite and not on the anterior margin of the paratergite
itself.
In the male genitalia, the sclerotized curved structure in the base of the
conjunctiva probably acts as a guiding mechanism for the vesica, which passes
along the ventral surface of this sclerite; the vesica is then directed on to the
V-shaped structure in the ventral wall of the conjunctiva and is thus everted.
PRUTHI (1925), in his account of the male genitalia of Chiloxanthus pilosus
(Fall.) and Salda littoralis (L.) did not mention the conjunctival appendages, the
ejaculatory reservoir, and the capitate processes. The latter, according to MARKS
(1951), are secondary developments in the Heteroptera, and mark the original
points of attachment of the parameres to the basal plates. EKBLOM (1926) also
failed to notice the ejaculatory reservoir and the capitate processes.
PRUTHI (1925) mentioned that the aedeagus in Mesovelia is not differentiated
into phallosoma and endosoma, but examination of Mesovelia mulsanti has shown
that the phallosoma, conjunctiva and vesica are distinguishable. EKBLOM (1928)
figured a drawn-out endosoma with its two appendages in his account of Meso-
velia furcata.
COMPARISONS
Head
On comparing the head structures of the two families, it is found that Saldula
and Mesovelia resemble each other in a number of features namely, the well
A. P. Gupta: Morphology of Saldidae and Mesoveliidae 189
developed vertex, the indistinct coronal and frontal ecdysial cleavage lines in
the adult, the well developed clypeal regions, the absence of the frontoclypeal
sulci, the four-jointed labrum, the antennae, and in the possession of four pairs of
cephalic trichobothria in the nymphs. However, these similarities lay not in
characters of great taxonomic importance in the group. There are a number of
important features in which they are quite distinct (Table II). It should be
mentioned that although Aepophilus bonnairei appears to resemble Mesovelia
more than Sa/dula, in respect of the most important taxonomic characters (that is,
the shape of the mandibular lever, the presence of epicranial pits, and the absence
of epipharyngeal process), Aepophilus is exactly like Saldula. It is evident that the
two families, the Saldidae and Mesoveliidae, show more differences than resem-
blances in their head structures, and thus it is concluded that in the head structure
they are distinct morphologically. The epicranial pits are a feature which appears to
be confined to the Saldidae and can thus be regarded as a character by which this
family can be separated from all other Heteroptera. However, other genera must
be examined to ascertain that these structures are actually a constant feature in
the family.
Thorax
In respect to the thoracic structures we also find that the Saldidae and
the Mesoveliidae show certain resemblances, particularly in the pronotal
collar, callal area with its median or lateral depressions, precoxal shelf,
trochantin, and fused metascutum and scutellum, well developed coxae, flattened
femora, and three tarsomeres, etc., but they exhibit distinct differences (Table II)
which outweigh the resemblances. A preliminary examination of some
of the families of the four higher taxonomic groups of the Heteroptera, namely
Pentatomomorpha, Cimicomorpha, Amphibicorisae and Hydrocorisae, revealed
that the structure of the mesonotum and the metasternum in the various families
are of either Saldula-type (the mesosternum projecting over the metasternum) or
Mesovelia-type (the mesosternum not projecting over the metasternum). Examin-
ation also revealed that, with the exception of the Amphibicorisae, the three
groups possess both types (GUPTA, in press). This indicates that if the above
mentioned taxa are natural groups, the structure of the thorax is of little value in
distinguishing the suprafamilial categories.
Abdomen
On comparing the structure of the female genitalia, the aedeagus, the presence
of ejaculatory reservoir, paratergites, and the seven pairs of abdominal spiracles,
the two families are found to be very similar, but it is evident that other
characters indicate that they are taxonomically distinct (Table II).
From the foregoing study it is evident that the Saldidae and the Mesoveliidae
are not closely related as suggested by the comparative morphological study of the
female genitalia; they are quite distinct in other morphological features.
190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
COMPARISON OF ALATE AND APTEROUS (AND BRACHYPTEROUS) FORMS IN THE
SALDIDAE AND MESOVELIJDAE
From the comparative morphology of the alate and the apterous forms in both
the Saldidae and the Mesoveliidae, it was found that these two groups show
distinct structural differences, particularly in the thorax (Table III). Insects with
flight possess well developed flight muscles and correlated with them an elaborate
thoracic structure, while those with limited or no flight have reduced flight mus-
cles, and consequently less developed thoracic structure. Flight in insects is effected
by two sets of muscles, the direct and the indirect. The indirect muscles include
the dorsal longitudinal muscles and the dorso-ventral muscles. The direct muscles
are attached to wing bases or wing sclerites and include principally the basalar,
the subalar and the muscles of the axillaries.
According to LARSEN (1945), the principal muscles are present in Saldula, and
accordingly, as one would expect, the apodemes and the internal margin of the
pronotal collar are well developed in the prothorax. The longitudinal muscles,
running from the first phragma to the second phragma in the pterothorax, mainly
produce the arching of the nota, and thus raising the notal processes relative to
the pleural processes, act as depressors of the wing. And since these muscles are
important in flight, the phragmata in Saldula are well developed. Similarly the
development of the furca seems to be correlated with the development of the
direct muscle, m. furca-pleuralis (of LARSEN). By the same token, the absence
of another direct muscle, m. coxa-subalaris (of LARSEN) both in the
mesothorax and the metathorax is correlated with the absence of the
subalar sclerites in the pterothorax. The lateral oblique muscle, m. mesonoti
secundus (of LARSEN) is well developed in the mesothorax of Saldula, but is
absent in the metathorax. This can be explained on the basis of WEBER's thesis
that in the Heteroptera the fore wings are principal organs of flight, and thus the
mesothorax is more developed than the metathorax. It is evident from the fore-
going that the morphological differences are the reflections of the functional
differences in the alate and the apterous forms. Unfortunately, no account of the
musculature of Mesovelia is available for comparison, but the structural dif-
ferences in the thorax of Mesovelia vittigera and M. mulsanti could also be
explained on a functional basis. This study has also revealed that the sternal region
in the alate and the apterous forms shows very little difference, and thus perhaps
the dorso-ventral muscles are not of great importance in the flight of these insects.
On comparing the alate and the apterous forms of both families, it is found
that the ocelli are rudimentary or absent in the apterous forms, but are present in
the alate forms. Accompanied with this presence of the ocelli in the alate forms,
are well developed compound eyes, which are not so conspicuous in the apterous
forms. Such correlated presence or absence of certain structures has also been
reported in the Lygaeidae (SCUDDER, personal communication). It is likely that
perhaps an alate insect needs more perfect visual apparatus than an apterous one,
and probably the ocelli supplement the compound eyes in their visual perception.
It is also possible that the presence of the ocelli in the alate forms and its absence
in the apterous may be due in part to a genetic linkage with some other
A. P. Gupta: Morphology of Saldidae and Mesoveliidae
character affected by the loss of flight.
The systematic position of the Saldidae and the Mesoveliidae, based on their
morphology, has been considered elsewhere (GUPTA, in press).
191
TABLE II. DIFFERENCES IN THE STRUCTURES OF HEAD, THORAX AND ABDOMEN
Parts Saldidae Mesoveliidae
HEAD
(1) Postclypeus. (1) Cephalic portion of (1) No such unpigmented
postclypeus with two areas present.
lateral unpigmented
areas (absent in
nymph): absent in
Aepophilus
(2) Paraclypeal region (2) Well defined in Sal- (2) Not well defined.
dula; not so in Aepo-
philus.
(3) Bucculae. (3) Well developed in (3) Not well developed.
Saldula; not so in
Aepophilus.
(4) Postgenal bridge. (4) Short in Saldula; lon- (4) Longer than in Sal-
ger in Aepophilus. dula.
(5) Mandibular lever. (5) Triangular both in (5) Roughly quadrangular.
Saldula and Aepophi-
lus.
(6) Postocciput. (6) Differentiated into dor- (6) Not so differentiated.
sal and lateral portions
in Saldula; not so in
Aepophilus.
(7) Maxillary plate area. (7) Not developed dorsal- (7) Developed dorsally.
ly in Saldula; develop-
ed dorsally in Aepo-
philus.
(8) Labrum. (8) Without epipharyngeal (8) With epipharyngeal
process both in Saldula process.
and Aepophilus.
(9) Cephalic trichobothria. (9) Eight to ten pairs in (9) Three pairs in the
adult Saldula; four adult.
pairs in Aepophilus.
(10) Compound eyes. (10) Conspicuous in Sal- (10) Not so conspicuous as
dula not so in Aepo- in Saldula.
philus.
(11) Ocelli. (11) Present in Saldula; ab- (11) Rudimentary in Meso-
sent in Aepophilus. velia mulsanti, pre-
sent in M. vittigera.
(12) Epicranial pits. (12) Present both in Saldu- (12) Absent.
la and Aepophilus.
THORAX
(13) Scutellum (mesoth.). (13) Extends over second (13) Does not extend over
abdominal segment in abdomen.
Saldula; not so in
Aepophilus.
(14) Parascutellum (meso- (14) Present in Saldula; ab- (14) Absent.
thorax). sent in Aepophilus.
(15) Pleural sulcus (meso- (15) Distinct but short in (15) Indistinguishable in
thorax).
both.
both.
(29)
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
Epimeron (mesoth.).
Sternum (mesoth.).
Coxal cleft (metatho-
rax).
Pleural sulcus (meta-
thorax).
Orifice of scent appara-
tus.
Fore wings.
Hind wings.
Base of media.
Tibial comb.
Pseudo-aroliae (par-
empodium).
Apodeme in sternum
VII.
Clasping organ.
Aedeagus.
Base of phallosoma.
Conjunctiva.
Ejaculatory reservoir.
Parameres.
(16)
(17)
(18)
(19)
(20)
(21)
(29)
(30)
(31)
(32)
Divided into dorsal
anepimeron and ven-
tral katepimeron.
Sternellum extending
over metasternum in
both.
Present in both.
Present and dorsal in
both.
Two, located laterally.
Differentiated into cla-
vus, embolium and
membranous part.
With jugal fold.
Not fused to costa,
subcosta and radius.
Absent in both.
Absent.
ABDOMEN
Absent.
Present.
Differentiated into
phallosoma, conjunctiva
and vesica in Saldula;
in Aepophilus, into
phallosoma and endo-
soma only.
Supported by an inver-
ted Y-shaped structure
in Saldula; by L-shap-
ed structure in Aepo-
philus.
Dorsal half sclerotized
into a curved structure
in both.
One.
Sickle-shaped.
(16)
(17)
(18)
(19)
(
29)
Not so divided in
both.
Sternellum not extend-
ing over metasternum.
Absent in both.
Absent in both.
One, median in posi-
tion.
Mostly membranous.
Without jugal fold.
Fused to costa, sub-
costa and radius.
Fore and middle tibiae
with combs at distal
ends.
Present.
Present.
Absent.
Differentiated into
phallosoma, conjunctiva
and vesica.
Absent.
Present.
Two.
Hook-like.
A. P. Gupta: Morphology of Saldidae and Mesoveliidae
193
TABLE II. DIFFERENCES IN THE ALATE AND APTEROUS (BRACHYPTEROUS) FORMS
IN THE SALDIDAE AND THE MESOVELIIDAE
Parts
Alate form
Saldula palustris
Apterous (brachypterous)
form
Aepophilus bonnairei
(1) Callal area. (1) Present. (1) Absent.
(2) Furcal arms. (2) Well developed. (2) Not well developed.
(3) Mesoscutellum. (3) Extends over the abdo- (3) Does not extend over
men. the abdomen.
(4) Ocelli. (4) Present. (4) Absent.
Mesovelia vittigera. Mesovelia mulsanti.
(CO) Callalkarear (1) Present. (1) Rudimentary.
(2) Mesonotum. (2) Differentiated into meso- (2) Mesonotum undifferen-
scutum and scutellum, tiated.
the former being over-
lapped by the posterior
part of the pronotum.
(3) Metanotum. (3) Median part extending (3) Does not extend over
over abdomen. abdomen.
(4) Ocelli. (4) Present. (4) Rudimentary.
KEY TO LETTERING OF FIGURES
A — anal vein EDS APP —
AC — anteclypeus EJD —
AEM — anepimeron EJR —
AF — antafossae 1 EM —
ANP — anterior notal wing process 2 EM —
AP — apodemes 3 EM —
APP — apophyseal pits EPP —
AS — antennal tubercle 1.ES —
ANAT — antenna 2, ES —
1 AX — first axillary 3 ES =
DAS — second axillary FC —
SPAN — third axillary FL —
B — spermathecal bulb FM —
BP — basal plates FR cs
1 BS — basisternum of prothorax FU —
2 BS — 5 „ mesothorax FWP —
3 BS — i ,, metathorax G —
BU — bucculae GA —
C — costa GP —
CA — callus
CE — compound eye GPC —
CL — clypeus GPL —
CNJ — conjunctiva 1 GPO —
CNJ APP — conjunctival appendages D (GEO ==
CO — corium 1 GX —
Ge — capitate processes DIE =
GI — coxal cleft ED —
CU — cubitus HWP —
CV — clavus J —
CX — coxa JE --
DMP — distal median process KEM —
EP — epicranial pit u —
EDS — endosoma LB —
endosomal appendages
ejaculatory duct
ejaculatory reservoir
epimeron of prothorax
a „ mesothorax
È „ metathorax
epipharyngeal process
episternum of prothorax
AR „ mesothorax
di „ metathorax
fecundation canal
flange of pump
femur
frons
furca
fore wing process
gena
gonangulum
- granular plate of clasping
organ
concavity of clasping organ
gonoplac
first gonapophysis
second gonapophysis
first gonocoxa
second gonocoxa
humeral plate
hind wing process
jugal vein
jugal fold
katepimeron
labium
labrum
194 TIJDSCHRIF VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
M — media PS — pleural sulcus
ME — membrane PSCU — parascutellum
MP — median plate 2 PSE — postscutellum of mesothorax
MSN — mesonotum 3, 1P8IE — 3 „ metathorax
MTN — metanotum PSN — postnotum
MXPA — maxillary plate area Pal — paratergite
N — notum PTAR — pretarsus
O — ocellus R — radius
OC — occiput SC — subcosta
OCC — occipital condyle DISC — mesoscutellum
OF — occipital foramen 3 SQL — metascutellum
PAC — paraclypeus 2 SCU — mesoscutum
PC — postclypeus 3 SCU — metascutum
PCS — precoxal shelf SGO — scent gland orifice
PG — postgena SPCD — spermathecal duct
PGB — postgenal bridge ST — stigma
2 PH — phragma of mesothorax 1 STL — sternellum of prothorax
SPF — Ri „ metathorax 2, SINE — „ mesothorax
PHS — phallosoma T — tergum
PHS APP — phallosomal appendages TAR — tarsus
PLA — pleurodema TB — tibia
PMP — proximal median plate. TN — trochantin
PN — pronotum INR — trochanter
PNC — pronotal collar i YY — first vannal vein
PNP — posterior notal wing process DY — second vannal vein
PO — postocciput VE — vannal fold
PR — paramere VG — vagina
1 PRS — presternum of prothorax VS — vesica
2 PRS — = „ metathorax VX — vertex
PRSC — prescutum WP — pleural wing process
V. ACKNOWLEDGMENTS
A part of this work was done while in receipt of a research grant from the
National Research Council of Canada, in the Department of Zoology, University
of British Columbia, Canada. I am thankful to Dr. G. G. E. SCUDDER for his
help during the progress of the work. Dr. C. J. DRAKE of the United States Na-
tional Museum kindly identified the species of Saldula. I am grateful to Dr. W. F.
BARR and Mr. A. R. Gittins for reading the manuscript and giving me the
benefit of their criticisms.
Furthermore, I am indebted to the Nederlandsche Entomologische Ver-
ceniging (Netherlands Entomological Society) which kindly accepted this paper
for publication.
VI. SUMMARY
The systematic position of the Saldidae is controversial. SCUDDER (1959) sug-
gested a close relationship between the Saldidae and the Mesoveliidae, based on
his studies of their female genitalia. The morphology of these two families has
been studied to throw some light on their probable relationship. It was found
that these two families are distinct morphologically. Their systematic position has
been considered elsewhere (GUPTA, in press).
A. P. Gupta: Morphology of Saldidae and Mesoveliidae 195
VII. LITERATURE CITED
BRINDLEY, M. D. H. 1934. The metasternum and pleuron of Heteroptera. Trans. R. ent.
Soc. Lond., vol. 82, p. 43—50.
COBBEN, R. H. 1960. Die Uferwanzen Europas, Hemiptera-Heteroptera, Saldidae. (In)
Wolfgang Stichel, Illustrierte Bestimmungstabellen der Wanzen II Europa, vol. 3,
p. 209—263.
DRAKE, C. J. and CHAPMAN, H. C. 1958. The sub-family Saldoidinae (Hemiptera-Saldidae).
Ann. ent. Soc. Amer., vol. 51, p. 480—485.
Drake, C. J. and Davis, N. T. 1958. The morphology and systematics of the Piesmatidae
(Hemiptera) with keys to world genera and American species. Ibid., vol. 51, p.
567—581.
DRAKE, C. J. and HoTTES, F. C. 1951. Stridulatory organs in Saldidae (Hemiptera). Great
Basin Nat., p. 11—43.
DuPorte, E. M. 1946. Observations on the morphology of the face in insects. J. Morphol.,
vol. 79, p. 371—418.
EKBLOM, T. 1926. Morphological and biological studies of the Swedish families of Hemi-
ptera-Heteroptera. Part I. The families Saldidae, Nabidae, Lygaeidae, Hydrometri-
dae, Veliidae and Gerridae. Zool. Bidr., Uppsala, vol. 10, p. 31—179.
EKBLOM, T. 1928. Morphological and biological studies of the Swedish families of Hemi-
ptera-Heteroptera. Part II. The families Mesoveliidae, Corizidae and Corixidae.
Ibid., vol. 12, p. 113—150.
FERRIS, G. F. 1944. On certain evolutionary tendencies in the heads of insects. Microento-
mology, vol. 9, p. 78—84.
GupTA, A. P. 1963. A consideration of the systematic position of the Saldidae and Meso-
veliidae. (In press, in Proc. Ent. Soc. Wash.).
Hoke, S. 1926. Preliminary paper on the wing venation of Hemiptera. Ann. ent. Soc. Amer.,
vol. 19, p. 13—34.
LARSEN, O. 1945. Der Thorax der Heteroptera, Skelett und Muskulatur. Lunds Univ.
Arsskrift. N.F. vol. 2, p. 41, 1945.
LESTON, D. 1957. The stridulatory mechanism in terrestrial species of Hemiptera-Hetero-
ptera. Proc. zool. Soc. Lond., vol. 128, p. 369—386.
MACGILI, E. I. 1947. The anatomy of head and mouthparts of Dysdercus intermedius Dist.
Proc. zool. Soc. Lond., vol. 177, p. 115—128.
Marks, E. P. 1951. Comparative studies of male genitalia of Hemiptera (Homoptera-Hete-
roptera). J. Kansas ent. Soc., vol. 24, p. 134—141.
Marks, E. P. 1959. The food pump of Pelocoris and comparative studies on other aquatic
Hemiptera. Psyche, vol. 64, p. 123—134.
Muir, F. and KERSHAW, J. C. 1911. On the homologies and mechanism of the mouthparts
of Hemiptera. Ibid., vol. 18, p. 1.
Murr, F. and KERSHAW, J. C. 1912. The development of the mouthparts in the Homoptera,
with observations on the embryo of Siphanta. Ibid., vol. 19, p. 77.
NEERING, T. 1945. Morphological variations in Mesovelia mulsanti (Hemiptera-Mesoveli-
idae). Univ. Kansas Bull., vol. 36, p. 126—148.
PARSONS, M. C. 1959. Skeleton and musculature of the head of Gelastocoris oculatus (Fabr.)
Hemiptera-Heteroptera. Bull. Mus. Comp. Zool., Harvard College, p. 122.
PENDERGRAST, J. G. 1957. Studies on the reproductive organs of the Heteroptera with a
consideration of their bearing on classification. Trans. R. ent. Soc. Lond., vol. 109,
p. 1—63.
PRUTHI, H. S. 1925. The morphology of the male genitalia in Rhynchota. Trans. R. ent.
Soc. Lond., 1925, pp. 121—267
SCUDDER, G. G. E. 1959. The female genitalia of the Heteroptera: morphology and bearing
on classification. Trans. R. ent. Soc. Lond., vol. 111, p. 405.
SMITH, J. B. 1892. Notes on the homology of the Homopterous mouth. Proc. Amer. Assoc.
Adv. Sci., vol. 40, p. 325
SNODGRASS, R. E. 1935. Principles of Insect Morphology. McGraw-Hill Book Company. Inc.
IND
196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 3, 1963
SNODGRASS, R. E. 1960. Facts and theories concerning the insect head. Smithsonian Misc.
Coll., vol. 142.
SPOONER, C. S. 1938. The phylogeny of Hemiptera based on a study of the head capsule.
Univ. Illinois Bull., no. 35, 70, p. 1—102.
TAYLOR, L. R. 1918. The thoracic sclerites of Hemiptera and Heteroptera. Ann. ent. Soc.
Amer., vol. 11, p. 225—249.
WEBER, H., 1929. Kopf und Thorax von Psylla mali. Zeitschr. Morph. Oekol., vol. 14, p.
59—165.
WEBER, H. 1930. Biologie der Hemipteren. Berlin.
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ED — 1568.2
i
DEEL 106 AFLEVERING 4 | JUI
TIJDSCHRIFT
VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
DE NEDERLANDSCHE ENTOMOLOGISCHE VEREENIGING
INHOUD:
R. STRAATMAN. — Observations on migration in certain Australian Lepidoptera,
Dor WSE
Tijdschrift voor Entomologie, deel 106, afl. 4. Gepubliceerd 15-V-1963
Nederlandsche Entomologische Vereeniging
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pu A rel as end odin
on
OBSERVATIONS ON MIGRATION IN CERTAIN
AUSTRALIAN LEPIDOPTERA
BY
R. STRAATMAN
At the time, Division of Entomology, C.S.I.R.O., Canberra, Australia
In Australia several species of Lepidoptera are known to have migratory habits.
WILLIAMS (1930) lists the following.
Catopsilia pomona pomona Fabr. Terias smilax Donovan
C. crocale Cramer Danaus plexippus L.
C. pyranthe pythias Waterhouse & D. chrysippus petilia Stoll.
Lyell D. hamata hamata Macleay
Appias paulina ega Boisduval D. affinis affinis Fabr.
Anaphaeis java teutonia Fabr. Acraea andromacha Fabr.
Delias nigrina Fabr. Vanessa cardui kershawi McCoy
D. harpalyce Donovan Badamia exclamationis Fabr.
The author had an opportunity to make a few observations on migration of
Lepidoptera which may prove to be of some value for other students. These
observations were made in the area between Ingham and Tully in north-eastern
Queensland (17—18° S.L.) in the years 1960—1961. The rainy season in
those years was unusually dry.
Towards the middle of March, 1961, Badamia exclamationis Fabr., a Hesperid
well known for its migratory habits, appeared in increasing numbers, flying con-
stantly in a north-north-westerly direction. The migration reached its peak in the
second half of April; a rough count yielded a total of 85—100 specimens per minute
crossing a stretch of approximately 80 yards of the main road, about one mile
from the sea shore between Ingham and Tully. Very few were seen feeding at this
site, but on the same day and again later in the month, in areas near rain forests
overgrown with Lantana many thousands of specimens were seen feeding at
flowers or drinking from moist sandy river banks, while many others were resting
on the underside of leaves. At Ingham several specimens were caught in a light
trap, suggesting that they were still active after nightfall. By the end of April their
numbers had decreased considerably, and from then on, only odd specimens were
seen, feeding actively but not migrating. No specimens were seen mating or ovi-
positing and no larvae were found. Of a few specimens caught in June, females
were somewhat predominant. It is not known, whether these specimens, which
were slightly worn, were hibernating.
The butterfly Danaus hamata Macleay was also abundant in the same area of
1977
198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 4, 1963
north-eastern Queensland. This species has a wide distribution in the Indo-Malayan
and Australian regions where it is represented by several races. From mid-February
odd specimens were seen flying consistently in a north-north-westerly direction.
Their number increased slowly but steadily, and their flight reached a peak by
mid-April. Rough counts made in April yielded a total of 55 to 65 specimens per
minute passing over a stretch of about 70 yards of road near Ingham. On several
occasions specimens were also taken in a light trap. A few specimens of Euploea
eichhorni Staudinger and E. corinna corina Macleay, were included in these
flights.
At Forest Beach, 12 miles east of Ingham, these butterflies were seen flying
towards the land from high over the sea, but on reaching the shore they dropped
to near ground level where many were found resting on scrubs and mangroves.
Little or no feeding was observed. At Mission Beach, 17 miles northeast of Tully
where rain forest grows near the beach, thousands of specimens were found in
the late afternoon clustering on small twigs, generally in clearings of the rain
forest sheltered from the wind. In the morning large numbers were observed
feeding on Lantana flowers and others were gathered around puddles on the
road.
As the season advanced, the number of migrating specimens decreased slowly,
although as late as June odd specimens were still flying in a north-north-westerly
direction. When migration was in progress, neither mating nor oviposition was
observed nor were any larvae found.
The first report about migratory flights of Australian butterflies comes from
Captain Cook, who mentions that at Thirsty Sound, on the East Coast of New
South Wales, he found an incredible number of butterflies “so that for the space
of three or four acres the air was so crowded with them that millions were to be
seen in every direction at the same time that every branch and twig was covered
with others that were not upon the wing”. In “Survey of the Coasts of Australia”,
Captain KING reports that at Cape Cleveland as well as at other places where he
landed, “the air was crowded with a species of butterfly, a great many of which
were taken”, and which he thought to belong to the same species recorded by
Captain Cook in Thirsty Sound. He further reports that this species was described
by Macleay as Euploea hamata. (Both reports are mentioned by MCKEOWN, 1944).
In the second half of June, at Forest Beach near Ingham, many thousands of
Danaid butterflies were found clustering on dry branches of the “paper-bark tree”,
Melaleuca leucodendron, which grows in and around swamps, about half a mile
from the sea shore. The dry conditions had caused the swamps to dry up, making
observations easy. Other specimens were found sheltering on the underside of dry
Pandanus and other leaves. Most specimens were Danaus hamata, the majority of
which were females, but there was a small percentage of two other species: Euploea
eichhorni Staud. and Hypolimnas bolina nerina Fabr. During the warmer hours of
the day many butterflies could be disturbed by walking beneath the trees, but Hypo-
limnas bolina remained inactive resting head downwards and antennae folded
between the wings. As soon as the disturbed Danaidae started to fly they were
attacked by birds, identified as the masked wood-swallow (Artamus personatus),
which caught the butterflies in a swooping flight, consuming the bodies but drop-
R. STRAATMAN : Migration of Australian Lepidoptera 199
ping the wings. Large quantities of wings on the ground suggested that many
butterflies had been destroyed by the birds. When a bird missed its prey, the
butterfly dropped to the ground, usually remained motionless for several minutes
and then escaped by slowly walking along the ground to take shelter in grass.
Those which flew up again were often caught. FRYER (1913) and WILLIAMS
(1927) recorded similar bird attacks on migrating Danaus hamata septentrionts
and on Euploea species, by the drongo, Dicrurus lencopygialis and by a wood-
swallow Artamus fuscus, in Ceylon. A well-known theory suggests that the
representatives of the family Danaidae which usually are slow fliers, besides being
conspicuously marked, would be distasteful to birds. It seems likely
now that Danaidae are distasteful only to some species of birds or that
they lose their distasteful properties while hibernating. Other Lepidoptera were
collected in the same area by beating dry Pandanus leaves, which form dense,
tent-like shelters around the trunks. Two species of Nymphalidae, Hypolimnas
bolina nerina Fabr. and H. alimena lamina Fruhst. and one of Pieridae, Catopsilia
pomona Fabr., were found in small numbers scattered throughout the area.
When disturbed the Nymphalids were quite inactive, generally dropping to the
ground where they either remained motionless or sometimes flew a short distance.
Catopsilia butterflies also flew only briefly before taking cover again. In these
species both sexes were present. None was seen feeding or flying, unless disturbed,
and their inactivity suggests that they were hibernating.
LITERATURE
BROEKHUYSEN, G. J. and J. G. P., 1960. — Butterfly Migration over Johannesburg. South
Afric. Journ. of Science, vol. 56, p. 10.
FRYER, J. C. F., 1913. — Field Observations on the Enemies of Butterflies in Ceylon.
Proc. Zool. Soc. London, p. 613—619.
McKeown, K. C., 1944. — Insect Wonders of Australia, p. 68. Angus & Robertson Ltd.,
Sydney.
WATERHOUSE, G. A., 1932. — What Butterfly is that? Angus & Robertson Ltd., Sydney.
WILLIAMS, C. B., 1927. — A Study of Butterfly Migration in South India and Ceylon.
Trans. Ent. Soc. Lond., vol. 75.
WILLIAMS, C. B., 1930. — The Migration of Butterflies. Oliver and Boyd, Edinburgh.
WILLIAMS, C. B., 1958. — Insect Migration. The New Naturalist, p. 13.
i
Ti
ee
i ti si te
ENTOMOLOGISCHE BERICHTEN
È
| De Entomologische Berichten worden eveneens door de Nederlandsche Entomo-
logische Verceniging uitgegeven en zijn bestemd voor de publicatie van kortere
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ringen en van mededelingen van het Bestuur. Zij verschijnen twaalf maal per jaar
in een aflevering van 16 of meer bladzijden. Deze 12 afleveringen vormen samen
een deel.
TRICHOPTERORUM CATALOGUS
Ook de Trichopterorum Catalogus van F. C. J. Fischer wordt door de Neder-
landsche Entomologische Vereeniging voor rekening van de auteur uitgegeven.
De eerste drie delen verschenen in 1960, 1961 en 1962; deel IV zal vóór medio
1963 uitkomen. Er volgen daarna nog elf delen, in totaal dus 15. Alle bekende
recente en fossiele soorten van de gehele wereld met hun synoniemen worden er in
behandeld. Van alle genera worden de type-soorten aangewezen en de literatuur
zal van 1758 tot eind 1960 met de geografische verspreiding vermeld worden.
De prijs van deel I en II is f 39,—, van deel III f 45,—. Van alle delen zijn ook
éénzijdig bedrukte exemplaren beschikbaar. De prijsverhoging hiervoor bedraagt
E2550:
Alle zakelijke correspondentie betreffende de Vereeniging te richten aan de
Secretaris, W. Hellinga, Weesperzijde 2311, Amsterdam-O.
Alle correspondentie over de redactie van het Tijdschrift voor Entomologie te
richten aan de Redacteur, Dr. A. Diakonoff, Rijksmuseum van Natuurlijke
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Alle correspondentie over de redactie van de Entomologische Berichten te rich-
ten aan de Redacteur, B. J. Lempke, Oude IJselstraat 12111, Amsterdam-Z. 2.
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29, Bergen (N.H.), postgiro 188130, ten name van de Nederlandsche Entomolo-
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Alle correspondentie betreffende de Bibliotheek der Vereeniging te richten aan
de Bibliotheek, Zeeburgerdijk 21, Amsterdam-O.
NOTICE TO SUBSCRIBERS
All correspondence referring to subscriptions and exchange subscriptions of the
periodicals of the Netherlands Entomological Society should be addressed to the
Librarian, Zeeburgerdijk 21, Amsterdam-O,
NOTICE TO CONTRIBUTORS
Contributors will receive free of charge fifty reprints of their papers, joint authors have
vo divide this number between them at their discretion. Additional reprints may be ordered
when returning proofs ; they will be charged at about two Dutch cents per page.
Manuscripts should be written in Dutch, English, French, German or Italian. If they
contain descriptions of new genera, species, etc. they should be in one of the four last
mentioned languages: only when the descriptions form a minor part of the paper, the
manuscript may be written in Dutch, with the descriptions in one of these languages. Papers
in Dutch should contain a short summary in one of these four languages.
Manuscripts should be typewritten in double spacing on only one side of the paper, with
a margir of at least three cm at the left side of each sheet. Paragraphs should be indented.
Carbon copies cannot be accepted, as handling makes them illegible.
Captions for text figures and plates should be written on a separate sheet in double
spacing, numbered consecutively in arabic numerals ; the use of a, b, c, or any other sub-
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Drawings for reproduction should be on good paper in Indian ink, preferably at least
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of the Tijdschrift, or a portion thereof. Combinations of illustrations into groups are
preferable to separate illustrations since there is a minimum charge per block.
Names of genera and lower systematic categories, new terms and the like are to be under.
lined by the author in the manuscript by a single straight line. Any other directions as to
size or style of the type are given by the editors, not by the author. Italic type or spacing
to stress ordinary words or sentences is to be avoided. Dates should be spelled as follows :
either ”10.V.1948” or ”10 May, 1948”. Other use of latin numerals should be avoided, as
well as abbrev’ ‘ions in the text, save those generally accepted. Numbers from one to ten
occurring in the text should be written in full, one, two, three, etc. Titles must be kept
short. Footnotes sl.ould be kept at a minimum.
Authors will be charged with costs of extra corrections caused by their changing of the
text in the proofs.
Bibliography should not be given in footnotes but compiled in a list at the end of the
paper, styled as follows:
Mosely, M. E., 1932. "A revision of the European species of the genus Leuctra (Pleco-
ptera)”. Ann. Mag. Nat. Hist., ser. 10, vol. 10, p. 1—41, pl. 1—5, figs. 1—57.
Text references to this list might be made thus:
"Mosely (1932) says.” or (Mosely, 1932)”.
The editors reserve the right to adjust style to certain standards of uniformity.
Manuscripts and all communications concerning editorial matters should be sent to
Dr. A. DIAKONOFF, Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands.
DEEL 106 AFLEVERING 5
TIJDSCHRIFT
VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
DE NEDERLANDSCHE ENTOMOLOGISCHE VEREENIGING
INHOUD:
C. J. H. FRANSSEN & W. P. MANTEL. — Supplement op de Naamlijst van
de in Nederland aangetroffen Thysanoptera, pp. 201—204.
Tijdschrift voor Entomologie, deel 106, afl. 5. Gepubliceerd 15-V-1963 |
Nederlandsche Entomologische Vereeniging
BESTUUR
Prof. Dr. J. van der Vecht, President (1961— 1967), Oegstgeest.
Dr. J. G. Betrem, Vice-President (1959—1965), Deventer.
W. Hellinga, Secretaris (1963—1969), Amsterdam.
Drs. H. Wiering, Penningmeester (1962—1968), Bergen (N.H.).
Drs. C. A. W. Jeekel, Bibliothecaris (1960—1966), Amsterdam.
F. C. J. Fischer (1958—1964), Rotterdam.
Dr. H. J. de Fluiter (1960—1965), Wageningen.
COMMISSIE VAN REDACTIE VOOR DE PUBLICATIES
Prof. Dr. J. van der Vecht (1961—1967), Oegstgeest.
P. Chrysanthus (1961—1964), Oosterhout (N.B.).
Dr. A. Diakonoff (1961—1964), Leiderdorp.
G. L. van Eyndhoven (1963— 1966), Haarlem.
Dr. L. G. E. Kalshoven (1961—1964), Blaricum.
Prof. Dr. D. J. Kuenen (benoemd 1957), Leiden.
Dr. P. A. van der Laan (benoemd 1957), Bennekom.
B. J. Lempke (1962—1965), Amsterdam.
Prof. Dr. J. de Wilde (benoemd 1957), Wageningen.
Drs. J. T. Wiebes (1963—1966), Leiden.
BESTUUR DER AFDELING VOOR TOEGEPASTE ENTOMOLOGIE
Dr. H. J. de Fluiter, Voorzitter, Wageningen.
Dr. F. J. Oppenoorth, Secretaris, Utrecht.
Mej. Dr. A. Post, Wilhelminadorp.
Dr. C. F. A. Bruyning, Oegstgeest.
Ir. P. Gruys, Rheden.
De contributie voor het lidmaatschap bedraagt f 15.—, voor student-leden
f 2.50, per jaar. — Begunstigers betalen jaarlijks tenminste f 15 —.
De leden, behalve de student-leden, ontvangen gratis de Entomologische
Berichten van 12 nummers per jaar, waarvan de prijs voor student-leden f 1.50
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De leden kunnen zich voor f 10.— per jaar abonneren op het Tijdschrift voor
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De eerder verschenen publicaties der Vereeniging zijn voor de leden tegen ver-
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TIJDSCHRIFT VOOR ENTOMOLOGIE
Het Tijdschrift voor Entomologie wordt uitgegeven door de Nederlandsche
Entomologische Vereeniging en is bestemd voor de publicatie van de resultaten
van de studie der Entomologie van algemene en bijzondere aard. Het verschijnt
in één deel van 300—400 bladzijden per jaar, bestaande uit enkele afleveringen.
Bovendien worden monographieën handelende over bijzondere onderwerpen,
met onregelmatige tussenpozen uitgegeven.
SUPPLEMENT OP DE NAAMLIJST VAN DE IN NE
AANGETROFFEN THYSANOPTERA
DOOR
C. J. H. FRANSSEN & W. P. MANTEL
Instituut voor Plantenziektenkundig Onderzoek, Wageningen
In 1962 publiceerden wij in dit tijdschrift een beredeneerde naamlijst van in
Nederland aangetroffen Thysanoptera. Intussen zijn er weer een aantal soorten
bijgekomen, waaronder één geheel nieuwe, die onlangs door TITSCHACK (1962)
beschreven werd. Veel van het ons ter beschikking staande materiaal werd ver-
zameld door de heer P. VAN UDEN, die daardoor een belangrijke bijdrage heeft
geleverd tot de kennis van de Nederlandse fauna.
Alvorens tot de behandeling van de nieuwe soorten over te gaan, moeten eerst
een drietal correcties in de oorspronkelijke lijst worden aangebracht. Als soort
no. 40 noemden wij Odontothrips loti Haliday; daarbij staat onder meer vermeld,
dat in de collectie DOEKSEN 17 9 en 2 4 werden aangetroffen onder de naam
Odontothrips ulicis Haliday. Wij hielden de betreffende dieren toen voor O. loti
Haliday, doch achteraf blijkt het door DOEKSEN verzamelde materiaal te behoren
tot de soort Odontothrips cytisi Morison, die nieuw is voor Nederland.
Onder de nummers 45 en 46 worden twee vertegenwoordigers van het ge-
slacht Rhopalandrothrips genoemd, namelijk consoctatus Targ.-Tozz. en obscurus
Targ.-Tozz. Laatst genoemde soort, die door Dr. H. PRIESNER op naam was ge-
bracht, is een synoniem van consociatns.
Tenslotte vermeldden wij onder no. 89 Haplothrips juncorum Bagnall. Deze
determinatie is niet juist geweest; een zekere determinatie is echter niet mogelijk
wegens het ontbreken van mannetjes.
Het aantal van 106 soorten in de naamlijst is hiermede teruggebracht tot 104,
doch met de 22 hieronder te behandelen nieuwe tripsen, zijn er thans 126 soorten
van Nederland bekend.
Veel dank zijn wij verschuldigd aan Prof. Dr. H. PRIESNER en Prof. Dr. E.
TITSCHACK voor het verifiëren van het door ons gedetermineerde materiaal.
De opgesomde soorten zijn macropteer, tenzij anders vermeld.
TEREBRANTIA
AEOLOTHRIPINAE
Aeolothrips ericae Bagnall
Maastricht: 1.VIII.1962, 4 9 op Genista pilosa L. en 1 © op Reseda lutea L.
Forma meulleri Priesner
Nunhem: 5.VI.1961, 3 ® op Vicia cracca L.; Haelen: 21.VII.1961, 7 2 op
Sarothamnus scoparius (L.) Wimm. (leg. P. van UDEN); Kessel-Eik: 22.V.1962,
1 9 op Cerastium arvense L. (leg. P. VAN UDEN) en 3 © op Sarothamnus sco-
201
202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 5, 1963
parius (L.) Wimm.; 29.V.1962, 5 © op Sarothamnus scoparius (L.) Wimm.;
Neer: 29.V.1962, 2 9 op Sarothamnus scoparius (L.) Wimm. (leg. P. VAN
UDEN).
Aeolothrips manteli Titschack
In totaal werden 68 stuks en 1 larve verzameld in de Amsterdamse waterleiding-
duinen te Vogelenzang (gem. Bloemendaal). Alle dieren werden gevonden op
Anchusa officinalis L. met uitzondering van 1 9, dat werd aangetroffen op een
V erbascum-soort. Het mannetje is nog niet bekend (FRANSSEN & MANTEL, 1963).
Aeolothrips propinquus Bagnall
Natuurreservaat „de Hamert’: 30.VII.1962, 1 9 op Echium vulgare L.; Maas-
tricht: 1.VIII.1962, 1 9 op Echium vulgare L.
Anaphothripina
Aptinothrips elegans Priesner
Deze aptere soort werd gevonden in de Amsterdamse waterleidingduinen te
Vogelenzang (gem. Bloemendaal). Het mannetje is nog niet van Nederland be-
kend (FRANSSEN & MANTEL, 1963).
Thripina
Iridothrips mariae Pelikan
Al het materiaal werd verzameld te Venlo op Typha latifolia L. in 1962: 22.V,
3,9, DOM, 3. Os ZEW Si Oven 3 8 MT N 2
Odontothrips cytisi Morison
Wageningen: 9.V.1937, 17 9 en 2 & op Sarothamnus vulgaris Wimm. (coll.
DOEKSEN); Kessel-Eik: 22.V.1962, 116 9 en 3 & op Sarothamnus scoparius (L.)
Wimm. en 1 2 op Betula-species; 29.V, 71 2 op Sarothamnus scoparius (L.)
Wimm.; Venlo: 22.V.1962, 1 9 op Typha latifolia L. (leg. P. VAN UDEN);
Tiel: 23.V.1962, 3 9 op Typha latifolia L. (leg. P. van UDEN); Neer: 29.V.
1962, 76 2 op Sarothamnus scoparius (L.) Wimm. (leg. P. VAN UDEN) en 1 9
op Sorbus aucuparia L. (leg. P. VAN UDEN).
Odontothrips meliloti Priesner
Heel en Panheel: 1.IX.1958, 2 9 op Meltlotus officinalis (L.) Lamk.; Venlo
10.VII.1962, 8 9 en 3 4 op Melilotus officinalis (L.) Lamk. (leg. P. VAN
UDEN).
Odontothrips phaleratus Haliday
St. Odiliënberg: 21.VII.1961, 4 9 en 1 4 op grassen.
Taeniothrips kratochvili Pelikan
Er werd één brachypteer wijfje verzameld te Wolvega (FRANSSEN & MANTEL,
1962).
C. J. H. FRANSSEN & W. P. MANTEL: Nederlandse Thysanoptera 203
Thrips alni Uzel
Al het materiaal van deze soort werd in 1962 verzameld te Kessel-Eik op Alnus
(edn aN(ES) sMoencht:)22:Ve SCO 2ON MSN O1 2 NIT 11179 Vents Tes"
Thrips minutissimus L.
Al ons materiaal behoort tot de forma obscura Coesfeld. Bloemendaal: 17.V.1961,
2 2 op Anchusa officinalis L. en 6.VI.1961, telkens één P op Anchusa of ficina-
lis L. en op grassen; Wolvega: 8.VI.1961, 1 9 op grassen (FRANSSEN & MANTEL,
1962 en 1963).
TUBULIFERA
Cryptothripina
Cryptothrips nigripes O. M. Reuter
Forma brachyptera
Haelen: 25.VI.1962, 1 9 en 3 & achter schors van Malws-species (leg. P. VAN
UDEN).
Forma macroptera
Haelen: 25.VI.1962, 4 9 achter schors van Malus-species (leg. P. VAN UDEN).
HAPLOTHRIPINI
Haplothrips-species
Wolvega: 22.VI.1961, 1.9 op grassen (FRANSSEN & MANTEL, 1962).
Neoheegeria verbasci Osborn
Haelen: op Verbascum-species, 21.VII.1961, 111 imagines en 425 larven; 14.VIII.
1961, 19 imagines, 149 larven, 22 voorpoppen en 109 poppen; Wellerlooi: 2.VII.
1962, 7 9 en 5 4 op Verbascum-species.
Xylaplothrips subterraneus J. C. Crawford
Wageningen: 25.XI.1960, 71 imagines op bollen van een Lilium-species.
Phlaeothripina
Hoplandrothrips bidens Bagnall
Kesteren: 23.VIII.1961, 1 2 op Symphoricarpos rivularis Suksd. (leg. P. VAN
UDEN).
Hoplandrothrips williamsianus Priesner
Kesteren: 23.VIII.1961, 2 2 op Symphoricarpos rivularis Suksd. (leg. P. VAN
UDEN).
Phlaeothrips annulipes O. M. Reuter
Wolvega: 25.V.1961, 1 ® op grassen (FRANSSEN & MANTEL, 1962).
204 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 5, 1963
Poecilothrips albopictus Uzel
Forma brachyptera
Wageningen: 9.VI.1962, 32 ® achter schors van Fagus sylvatica L. (leg. R. CoB-
BEN).
Forma macroptera
Wageningen: 9.VI.1962, 15 Q achter schors van Fagus sylvatica L. (leg. R. CoB-
BEN).
Cephalothripina
Cephalothrips monilicornis O. M. Reuter
Forma aptera
Bloemendaal: op grassen, 6.VI.1961, 9 9, 16.VI.1961, 29 en 1 larve, 12.VII.
1961, 2 imagines, 10.VIII.1961, 20 imagines, 8.VI.1962, 16 imagines.
Forma macroptera
Bloemendaal: 10.VIII.1961, 1 9 op grassen.
Hoplothripina
Hoplothrips longisetis Bagnall
Lichtenvoorde: 31.X.1961, 2 9 achter schors van Quercus-species (leg. P. VAN
UDEN); Aerdenhout: 26.1.1962, 1 ® achter schors van Quercus-species.
Hoplothrips ulmi F.
Forma brachyptera
Bleiswijk: 28.VIII.1961, 6 © en 2 & achter schors van Prunus avium L. (leg. P.
VAN UDEN).
Forma macroptera
Grollo: 10.VI.1961, 2.9 op grassen onder Ulmus carpinifolia Gled.; Bleiswijk:
28. VIII.1961, 5 9 achter schors van Prunus avium L. (leg. P. VAN UDEN).
SUMMARY
A supplement is given to the list of Thysanoptera (Tijdschrift voor Entomolo-
gie, 1962, vol. 105, p. 97—133). The material has been collected in 1961 and
1962 by Mr. P. vAN UDEN and both the authors of this paper. Up to now
126 species have been found in the Netherlands.
LITERATUUR
FRANSSEN, C. J. H. & MANTEL, W. P., 1962. Lijst van in Nederland aangetroffen Thysano-
ptera met beknopte aantekeningen over hun levenswijze en hun betekenis voor onze
cultuurgewassen. Tijdschrift voor Entomologie, vol. 105, p. 97—133.
————, 1962. Thysanoptera, in 1961 verzameld in de natuurreservaten „De Bennekomse
Meent” te Bennekom (Geld.) en „De Lindevallei” te Wolvega (Fr.). Natuur-
historisch Maandblad, vol. 51, p. 46—48.
— 2. 1962. Liothrips vaneeckei en Xylaplothrips subterraneus, twee voor lelies schadelijke
blaaspoten. Tijdschrift over Plantenziekten, vol. 68, p. 285—288.
—, 1963. Thysanoptera, verzameld in de Amsterdamse Waterleidingduinen. Natuur-
historisch Maandblad, vol. 52, p. 12—13.
TITSCHACK, E., 1962. Thysanoptera XXV. — Aeolothrips manteli nov. spec., ein neuer
Blasenfuss aus Holland. Bombus, vol. 2, p. 133—137.
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MUS. COMP. Z00L
LIBRARY
SEP 16 1963
UADGUAR
HARVARD
UNBERSITY
è | OSDL
DEEL 106 AFLEVERING 6
|
TIJDSCHRIFT
VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
DE NEDERLANDSCHE ENTOMOLOGISCHE VEREENIGING
SÌ
GE
U js
SEG
2
INHOUD:
C. A. W. JEEKEL. me oa toe from Borneo (Diplopoda, Polydesmida),
PPO 283-
Tijdschrift voor Entomologie, deel 106, afl. 6. Gepubliceerd 19-VIII-1963 |
Nederlandsche Entomologische Vereeniging
BESTUUR
Prof. Dr. J. van der Vecht, President (1961—1967), Oegstgeest.
Dr. J. G. Betrem, Vice-President (1959—1965), Deventer.
W. Hellinga, Secretaris (1963—1969), Amsterdam.
Drs. H. Wiering, Penningmeester (1962—1968), Bergen (N.H.).
Drs. C. A. W. Jeekel, Bibliothecaris (1960—1966), Amsterdam.
F. C. J. Fischer (1958—1964), Rotterdam.
Dr. H. J. de Fluiter (1960—1965), Wageningen.
COMMISSIE VAN REDACTIE VOOR DE PUBLICATIES
Prof. Dr. J. van der Vecht (1961—1967), Oegstgeest.
P. Chrysanthus (1961—1964), Oosterhout (N.B.).
Dr. A. Diakonoff (1961—1964), Leiderdorp.
G. L. van Eyndhoven (1963—1966), Haarlem.
Dr. L. G. E. Kalshoven (1961—1964), Blaricum.
Prof. Dr. D. J. Kuenen (benoemd 1957), Leiden.
Dr. P. A. van der Laan (benoemd 1957), Bennekom.
B. J. Lempke (1962—1965), Amsterdam.
Prof. Dr. J. de Wilde (benoemd 1957), Wageningen.
Dr. J. T. Wiebes (1963—1966), Leiden.
BESTUUR DER AFDELING VOOR TOEGEPASTE ENTOMOLOGIE
Dr. H. J. de Fluiter, Voorzitter, Wageningen.
Dr. F. J. Oppenoorth, Secretaris, Utrecht.
Mej. Dr. A. Post, Wilhelminadorp.
Dr. C. F. A. Bruyning, Oegstgeest.
Ir. P. Gruys, Rheden.
De contributie voor het lidmaatschap bedraagt f 15.—, voor student-leden
f 2.50, per jaar. — Begunstigers betalen jaarlijks tenminste f 15 —.
De leden, behalve de student-leden, ontvangen gratis de Entomologische
Berichten van 12 nummers per jaar, waarvan de prijs voor student-leden f 1.50
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TIJDSCHRIFT VOOR ENTOMOLOGIE
Het Tijdschrift voor Entomologie werdt uitgegeven door de Nederlandsche
Entomologische Vereeniging en is bestemd voor de publicatie van de resultaten
van de studie der Entomologie van algemene en bijzondere aard. Het verschijnt
in één deel van 300—400 bladzijden per jaar, bestaande uit enkele afleveringen.
Bovendien worden monographieën handelende over bijzondere onderwerpen,
met onregelmatige tussenpozen uitgegeven.
maten shal ou SL Lat m Sn Gend
pr
PARADOXOSOMATIDAE FROM BORNEO ~~
(DIPLOPODA, POLYDESMIDA)
BY
C. A. W. JEEKEL
Zoölogisch Museum, Amsterdam
INTRODUCTION
In comparison with the numbers of Paradoxosomatidae*) known to occur in
Java or Sumatra, remarkably few species of this family have been recorded from
Borneo. In all, the faunistic list of the island included the following species:
Orthomorpha beaumontii (Le Guillou) Orthomorpha kuekenthali (Att.)
Orthomorpha coarctata (Sauss.) Pratinus fasciatus (Att.)
Orthomorpha rotundata Att. Euphyodesmus gracilis Att.
Orthomorpha borneona Att. ‘Strongylosoma” nodulosum Att.
Of these species, O. beaumontii was described in 1841 in a way which rendered
its identity and relationship largely a matter of speculation. Similarly, "S.” nodu-
losum, because of having been based on a female specimen, always remained a
species of doubtful status. O. coarctata, finally, is a circumtropical ubiquist, which
probably does not belong to the autochthonous fauna of the island. In reality,
therefore, there remained but five species to show us the particularities of the
Paradoxosomatid fauna of Borneo.
That this small number of species represented a negligible portion of the local
fauna was revealed to me by the study of a rather large amount of material of
Paradoxosomatidae from Borneo which I found in various lots of unidentified
millipeds received in loan from the Museums of Bogor, Leiden, and Stockholm.
The collections under consideration cover the eastern, central and western parts
of Borneo and were made in particular by Dr. A. W. NIEUWENHUIS, in the course
of several Dutch Borneo Expeditions of some sixty years ago, by Dr. E. MJOBERG
during the middle twenties and, more recently, by Mr. A. M. R. WEGNER. For
the loan of these valuable collections I am very much indebted to the authorities
of the above mentioned Museums, and in particular to Miss A. M. BUITENDIJK 7
(Leiden), Mr. L. J. M. Buror (Bogor) and Dr. R. MALAISE (Stockholm).
My thanks are also due to Dr. M. VACHON (Paris) for enabling me the re-
examination of the type specimen of Orthomorpha beaumontn.
The present study not only multiplies the number of known species but also
shows that Borneo has a Paradoxosomatid fauna which, although having general
*) The name Paradoxosomatidae Daday, 1889, has priority over Strongylosomatidae,
Cook, 1895.
205
206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
southeast Asiatic features, is characterized by a large percentage of endemic genera.
In fact, the examination of the material under report has revealed the existence of
several generic categories not recognised previously. Especially the species hitherto
referred to Orthomorpha upon closer study have proved to belong to various groups
of questionable interrelationship. In order to arrive at a better taxonomy, it has
been deemed necessary to separate generically these groups from Orthomorpha.
The possibility that some of the proposed generic names in the future will
prove to have only subgeneric value has, of course, to be left open. For the present
it seems better to refrain from a more refined taxonomy in a group where so
many forms apparently await discovery.
For similar reasons I have abstained from using a trinary nomenclature. In
general, the decision by previous authors on what constitutes a subspecies appears
to have been influenced by the degree of morphological differentiation rather
than by genetic and geographical coherence. In the present paper several instances
can be found where morphologically very similar forms occur in one locality. Not
seldom the differences between such forms, to be looked upon as “good” species,
are less significant than those between two forms which may represent geogra-
phically vicarious populations. I have mentioned, as a rule, the possibility of sub-
specific relationship where necessary.
With the present additions the list of Paradoxosomatidae of Borneo reads as
follows:
pag. pag.
Kalimantanina gen. nov.. . . 207 cornalata spec.nov.. . . . 247
AIG IA SEC Nov 22208 GEENT AIS PECH Nov ES
hirttarsus spec. nov. » . . 212 pallıdasrspeenos eos
OGO alas SOs, Ms 025 OSC (LNG) eo 295)
alvonigrd spec nov NP PIS inchopleuna spec, none. a) a8 255
decolorata spec. nov. . . - 216 SDINENCENS SPEC OVA ans, 259
BOMmeoniia gen" OVEN meee Orihomorpha Boll) 2 77 277261
RHO Sores DON NT 9) 5 ZU beaumonti (Le Guill.) . . 269
DAA. O 8 | PAD) Goanelata, (Saussyy 02000200272
OC4/GIAR SPEC NOV 20223 Oxidas Cook an een eye
RODEO. (UNE), 226 kuekenthali (Att). . . . 273
Arthrogonopus gen. nov.. . . 227 JENIN OOO IS ING gw 275
edentulus Specs nova 2227228 Ord Ae INE ie van ereen 275
denticulatus spec. nova. N. 9 23 postfemoralis spec. nov. . . 279
DONT CON CALE MERE 25 granulifer spec. nov. . . . 280
CETUS SEE, WN 6) 6 ie be PRE “Strongylosoma” nodulosum
Gigantomorpha gen. nov. . . 237 EINECS ES A ate 2 Ol
LINMANIS SPEC NOV... SN ie 242
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 207
DESCRIPTIONS
Kalimantanina gen. nov.
Generic diagnosis. — 20 segments. Poreformula normal. Head without par-
ticulars; labrum tridentate. Antennae rather long.
Segments moderately (@ ) or rather strongly ( & ) constricted. Prosomites with
fine cellular structure, somewhat dull. Waist of moderate width, strongly long-
itudinally ribbed in the dorsal half of the segments, finely and rather widely
striate in the ventral half dorsad of the sternal level. Metatergites shining, hair-
less or with few hairs, without sculpture. Transverse furrow distinct but not very
deeply impressed, finely longitudinally striate, present from the 5th segment on-
wards. Pleural keels well developed in a number of segments in the anterior half
of the body.
Lateral keels rather weakly developed, posteriorly rounded on nearly all seg-
ments and projecting behind the posterior margin of the metasomite only in the
2nd segment, or, eventually, also very slightly in the 18th and 19th segments.
Keels of 2nd segment well below the level of those of the next segment.
Sternites with moderately to well developed cross-impressions, longer than wide
(4) or about as long as wide (?), rather weakly to moderately setiferous.
Sternite of 5th segment of & with a well developed, distally laminate process
between the anterior legs. Sternite of 6th segment of & somewhat modified:
only its anterior part somewhat raised above the ventral surface of the metasomal
ring. Sternite of 7th segment of & with on each side a latero-anterior sub-
longitudinal ridge-like protuberance. Sternite of 8th segment of 4 not modified.
Legs rather long; the distal part of the tibiae and the tarsi of at least some of
the anterior legs of the & with brushes.
Gonopods with coxa of moderate size, straight-cylindrical, with the distal end
slightly bent caudad; latero-anterior side of the distal end with a setiferous area.
Prefemur somewhat elongate, almost in one line with the femur and laterally
well demarcated from that joint, the demarcation oblique on the longitudinal axis
of the femur. Femur straight, well developed, about as long as coxa or somewhat
longer. Postfemur not demarcated. Spermal channel running more or less straight
along the anterior or medio-anterior side of the femur towards the base of the
solenomerite. Solenomerite long, flagelliform, arising from the anterior side of the
distal end of the femur. From the medio-anterior side of the femur, more or less
near the base of the solenomerite, or from the solenomerite itself, directly mesad
of the course of the spermal channel, arises a rather long, lanceolate or spiniform
process. Tibiotarsus springing from the posterior side of the distal end of the
femur, well developed, distinctly demarcated from the femur. Tibiotarsus con-
sisting of a simple, more or less circular solenophore without accessory processes,
curving into a posterior direction. Both lamina medialis and lamina lateralis well
developed, sheathing the solenomerite for its greater part.
Type-species. — Kalimantanina ruficeps spec. nov.
Range. — Borneo.
Number of species. — Five.
208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Remarks. — In a previous paper (JEEKEL, 1953), in connection with the
description of a new species, I have discussed already the heterogeneity within
the genus Sundanina Att. In particular I then questioned the dominant taxonomic
value previously attributed to the presence of one or more femoral processes in
the gonopods of the species of that genus. When the general structure of the
gonopods is taken into consideration, fundamental diversities are encountered
which seem to exclude the possibility that the species of Sundanina really represent
a monophyletic unity.
On account of the presence of a femoral process the species of Kalimantanina
could have been referred to Sundanina in the broad sense of ATTEMS. However,
pending a revision of the latter genus it seems more convenient to separate at
once the homogeneous group of presently described Bornean species. As a matter
of fact, there seems to exist no particular relationship between Kalimantanina and
any of the known species of Sundanina. The genus is, however, closely related to
Borneonina gen. nov.
Key to the species. — In general morphology the species of Kalimantanina are
extremely similar. For their separation we have to rely mainly on the characters
of the male gonopods.
1. Femoral process of gonopods arising well proximad of the base of the
solenomerite and projecting scarcely distad of the base of the tibiotarsus ...
Ree K. decolorata spec. nov.
— Femoral process arising from the base of the solenomerite, or from the
SOlEnOMEEEERPrOPEE NI OO EE ee re 2
2. Dorsum and sides of the somites black; head and anal valves of a contrasting
reddish: brownacolour TI 3
— Dorsum and sides of the somites black, except the lateral keels and an area
dorsad of these which are white. Head and anal valves not contrasting in
COLOUR ii CT E 4
3. Femur of gonopods about as long as the coxa. Femoral process arising from
the solenomerite. Tarsal brushes of the ¢ absent in the legs of the posterior
halfsofgthenbodyen nn. ee. K. ruficeps spec. nov.
— Femur about one and a half times as long as coxa. Femoral process arising
just proximad of the base of the solenomerite. Tarsal brushes of the & absent
Onlysinutheslast two) pairsioty legs eso. werner K. hirtitarsus spec. nov.
4. Femur of gonopods about as long as the coxa. Femoral process spiniform.
White area above the lateral keels about as broad as the keels ..................
Pe ene Rr IE RATTI K. albonigra spec. nov.
— Femur about one and a half times as long as coxa. Femoral process lanceolate.
White area dorsad of the lateral keels only half as wide as the keels .........
SIRO RAM TI OR NE EER Ea it ei o K. ocellata spec. nov.
Kalimantanina ruficeps spec. nov.
Material. — East Borneo: Pajan River (Coll. Dr. E. MJÖBERG, Mus. Stock-
holm) 25712702);
Colour. — Head reddish brown, very dark in the vertigial part. Antennae
brownish yellow, the 6th joint growing very dark, almost blackish brown, towards
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 209
the end. 7th joint also very dark, but the distal portion and the 8th joint whitish.
Collum and subsequent segments black, the poriferous keels very dark brown
around the pore. Ventral portion of somites brownish gray. Sternites and legs
dirty pale yellow. Anal segment black, the tail growing brownish yellow towards
the end. Valves and anal scale reddish brown.
Width. — Holotype 4 2.5 mm, other & 2.6 mm; 9 3.2 mm and 3.4 mm.
Head and antennae. — Labrum narrowly and moderately emarginate. Clypeus
rather weakly convex, moderately impressed towards the labrum. Lateral sides
hardly emarginate. Surface of head shining, with some irregular wrinkles. Clypeal
part of headplate rather dispersedly setiferous, a few bristles on the frontal part
and two on the vertex. Antennal sockets separated by slightly more than the dia-
meter of one socket or by about three fifths of the length of the 2nd antennal
joint. Postantennal groove shallow, the wall in front moderately prominent. Ver-
tex moderately convex, not demarcated from the frontal area. Sulcus moderately
impressed, with some transverse wrinkles, reaching downward to about the upper
level of the sockets. Antennae not particularly slender. Length of joints decreasing
very slightly from the 2nd to the Sth, the 6th joint somewhat more distinctly
shorter than the 5th, about two thirds of the length of the 2nd. Joints of subequal
width, the 6th very slightly thicker than the others. Antennal joints moderately
to rather densely setiferous.
Collum. — (fig. 1—2). Sligthly narrower than the head, subsemicircular in
dorsal outline. Anterior border evenly rounded, posterior border widely and rather
weakly emarginate. Lateral sides rather widely rounded. Surface smooth and
shining, longitudinally as well as transversely moderately convex; a few hairs
may be present. Lateral and latero-anterior border with a fine marginal rim,
gradually disappearing towards the middle of the anterior border.
Body segments. — Metatergites mostly hairless. Transverse furrow present up to
the 17th segment, extending laterad to about halfway the dorsal delimitation of
the lateral keels in most segments. Sides rugulose and dispersedly granulate up to
the 4th segment, only weakly rugulose from the 5th segment onwards. Pleural
keels present up to the 7th segment, totally absent from the 8th segment onwards.
Up to the 4th segment these keels are represented by complete ridges which are
posteriorly produced into a well developed angular lappet projecting behind the
posterior margin of the somites. From the 5th segment onwards the pleural keels
are present only on the posterior half of the segments and the posterior lappet
becomes smaller, projecting very slightly behind the margin of the somite in the
5th segment only.
Lateral keels. — (fig. 1—4). 2nd segment somewhat wider than the collum.
The keels anteriorly somewhat shouldered at the base, the anterior and posterior
edges rounded, the posterior edge projecting slightly behind the border of the
segment. Marginal rim rather weakly developed. 3rd and 4th segments of sub-
equal width, slightly narrower than the 2nd. The keels anteriorly widely, posterior-
ly somewhat more narrowly rounded, ventrally demarcated only in their posterior
half. 5th segment distinctly wider than the 4th. Keels from the 5th segment on-
wards anteriorly and posteriorly widely rounded, especially in the poreless seg-
ments. Poriferous keels somewhat more prominent. Only in the 18th and 19th
segments the keels have a minute acutely angular posterior edge, which, however,
210
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 211
does not project behind the margin of the segments. Dorsai furrow of keels
narrow, not reaching the waist. Seen from the lateral side the poreless keels have
their dorsal margin slightly concave, the poriferous keels are somewhat convex.
Poreless keels ventrally demarcated only near the posterior end, poriferous keels in
the posterior half. Pores lateral, in a slight excavation, somewhat more near the
ventral demarcation of the keels.
Sternites and legs. — Sternites of middle segments one and one third times as
long as broad, rather weakly setiferous. Cross impressions rather wide, not sharply
impressed. Sternite of 5th segment with a parallel-sided process directed cephalo-
ventrad and projecting slightly before the anterior border of the sternite. End of
process rounded and indistinctly bilobate by a very weak median incision, the
anterior side densely set with short bristles. Behind the process a rather deep
transverse furrow; between the posterior legs a normal longitudinal impression.
Sternite of 6th segment posteriorly not raised above the ventral level of the
metasomite; the anterior portion widely transversely concave and slightly longitu-
dinally convex. Legs (fig. 5) moderately to, in tarsi, rather densely setiferous
Tarsi and distal end of tibiae of the anterior legs with ventral brushes, which are
rapidly thinning out in the subsequent legs and are absent in the legs of the
second half of the body. Length of joints: 3>6>5>4 = 251.
Anal segment. — Tail rather broad at its base, dorsoventrally rather thick, of
moderate length. The sides rather strongly and straight converging; before the
end a well developed lateral setiferous tubercle, the end truncate and scarcely
emarginate. Anal segment with a very slight transverse depression at the base of
the tail. Ventral side of tail somewhat concave. Valves with narrow rims, the
setae on small tubercles. Scale triangular, the end truncate, the sides weakly
concave. The setiferous tubercles at the posterior edges rather close to each other,
projecting very weakly.
Gonopods. — (fig. 6). Femur of about the same length as the coxa. Femoral
process (p) spiniform, arising from the solenomerite. Tibiotarsus relatively well
developed, curving somewhat mesad and caudad, then proximad, and, finally,
somewhat laterad.
Female. — In the tube containing the males of ruficeps and the holotype
of ocellata there were two female specimens, obviously belonging to Kalimantanina
and with some doubt referable to the former of the two species. These two .9
differ from the 4 of ruficeps, aside from the usual secundary sexual characters
like the modifications of the sternites of the anterior segments, the brushes of
the legs, etc., in the following particulars. Colour either faded or not fully
developed: except for a very fine black middorsal stripe, entirely pale brownish yel-
low. The antennae have the distal half of the 6th and the proximal part of the
7th joint dark brown. Antennae somewhat more slender than in the ¢, the 6th
Fig. 1—6. Kalimantanina ruficeps spec. nov. — 1: left side of the head and the four
anterior segments of the holotype 4, lateral view. 2: left side of the head and the three
anterior segments of the holotype 4, dorsal view. 3: left side of the 10th and 11th segments
of the holotype 4, dorsal view. 4: the same, lateral view. 5: leg of the 7th segment of
the holotype 4. 6: right gonopod of the holotype 4, medial view. p: femoral process; 1:
lamina lateralis; m: lamina medialis. (The solenomerite has moved out of its natural
position)
212 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
joint being as wide as the proximal joints. Body segments comparatively less con-
stricted in the waist, resulting in a more robust aspect of the animals. Pleural keels
as in the 4, but missing from the 7th segment onwards. The lateral keels are
somewhat more prominent than in the 4; the posterior edges of the keels of
the 17th segment are minute but acutely angular. Sternites about as long as broad.
Legs scarcely more slender than in the ¢.
Kalimantanina hirtitarsus spec. nov.
Material. — East Borneo: without nearer location (Coll. Dr. E. MJÖBERG,
Mus. Stockholm), 1 ¢ (holotype).
Differing from rzficeps in the following characters.
Colour. — On the whole a little darker, probably because of a somewhat better
Fig. 7—8. Kalimantanina hirtitarsus spec. nov. — 7: tibia and tarsus of the leg of the 7th
segment of the holotype 4. 8: right gonopod of the holotype ¢, medial view
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 213
state of preservation. In the body segments only the venter is brownish gray. The
end of the tail dark brown.
Width. — 3.1 mm.
Head and antennae. — Vertigial sulcus running downward nearly to the lower
level of the antennal sockets.
Sternites and legs. — Legs (fig. 7) with tarsal and distal tibial brushes rather
dense, gradually thinning out in the legs of the second half of the body, but
absent only in the two ultimate pairs.
Gonopods. — (fig. 8). Femur about one and a half times as long as coxa.
Femoral process serrulate, arising from the base of the solenomerite. Tibiotarsus
relatively smaller than in rzfzceps, curving medio-distad, caudad and a little prox-
imad, and finally, latero-distad. Lamina medialis with an anterior emargination.
Female. — Unknown.
Remarks. — On the whole this species is so similar to ruftceps, that it might
as well be regarded as a geographical race of the latter. However, as compared to
the other species of the genus the differences in the gonopods of ruficeps and
hirtitarsus are quite significant, and a specific treatment seems fully justified.
Kalimantanina ocellata spec. nov.
Material. — East Borneo: Pajan River (Coll. Dr. E. MJOBERG, Mus. Stockholm),
1 g (holotype). East Borneo: without nearer location (Coll. Dr. E. MJOBERG,
Mus. Stockholm), 3 4,1:9.
Differing from r#ficeps in the following characters.
Colour. — Head very dark brown, with only the labral area somewhat paler.
Antennae dark castaneous, the distal part of the 7th and the 8th joints whitish.
Groundcolour of collum and body segments very dark brownish gray to black.
Lateral sides of collum and of the metasomites of the subsequent body segments
with rather large, rather sharply demarcated, white spots, comprising the lateral
keels and an area dorsad of these of about half the dorso-ventral width of a
poriferous keel. Medial side of these spots rounded. Ventral side of body segments
dark. Sternites yellowish; the legs brownish yellow, with the tarsi in the distal
half shading to dark brown. In a few pairs of anterior legs the dark colour
extends over the whole length of the tarsus and over the distal portion of the
tibia. Anal segment dark brownish gray to black, valves and scale paler brown.
Width. — Holotype 3.2 mm, paratypes & 3.0, 3.4 and 3.4 mm; allotype 9
3.8 mm. ;
Head and antennae. — Frontal area of head sometimes demarcated from vertex
by a weak depression. Vertigial sulcus running downward to about the upper
level of the antennal sockets, or continued below that level by a slight longitudinal
impression.
Collum. — Posterior border at each side dorsad of the lateral roundings with a
weak notch.
Body segments. — Pleural keels present up to the 7th segment, almost
completely absent on the 8th. The posterior lappets projecting behind the posterior
margins up to the 6th segment.
Lateral keels. — (fig. 9). Somewhat more prominent than in ruficeps or hir-
214 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
titarsus; the keels of the poreless segments have the posterior edges distinctly more
narrowly rounded, they are ventrally demarcated in their posterior half. Keels of
the 18th and 19th segments as in r#ficeps.
Sternites and legs. — Sternites one and a half times longer than broad. Cross-
impressions somewhat deeper. Sternal process of 5th segment with the sides weakly
converging in the distal direction. Brushes of tibiae and tarsi as in hirtitarsus:
absent only in the last two pairs of legs.
Anal segment. — Sides of tail somewhat less converging distally; the end
somewhat more distinctly emarginate.
Gonopods. — (fig. 10). Femur about one and a half times as long as coxa,
widening distad. Femoral process arising from the base of the solenomerite,
elongate, laminate and more or less curving in a spiral. Tibiotarsus curving medio-
distad, caudad and finally laterad and a little distad.
Female. — In the Sth to the 7th segments the pleural keels are somewhat less
strongly developed than in the 4. Sternites slightly longer than broad. The legs
comparatively a little shorter than in the 4.
Fig. 9—10. Kalimantanina ocellata spec. nov. — 9: left side of the 10th and 11th segments
of the holotype 4, dorsal view. 10: right gonopod of the holotype 4, medial view
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 215
Remarks. — The gonopods in the three male paratypes are almost identical
with those of the holotype. The telopodite, however, is slightly tortuous towards
the lateral side, so that the femoral process slightly projects caudad of the femur.
Kalimantanina albonigra spec. nov.
Material. — East Borneo: Mt. Tibang, 1700 m (Coll. Dr. E. MJÔBERG, Mus.
Stockholm), 1 4 (holotype). East Borneo: without nearer location (Coll. Dr. E.
MJÖBERG, Mus. Stockholm), 1 &.
Differing from ruficeps in the following characters.
Colour. — As in ocellata, but the white lateral spots of the collum and meta-
somites are comparatively larger and embrace the keels and an area dorsad of
these of about the dorso-ventral width of a poriferous keel. Legs paler: yellowish
white.
Width. — Holotype 2.9 mm, paratype 2.7 mm.
Head and antennae. — Vertigial sulcus running downward to just below the
upper level of the antennal sockets.
Collum. — Posterior border dorsad of the lateral roundings very weakly emar-
ginate.
Body segments. — Pleural keels as in ocellata.
Fig. 11. Kalimantanina albonigra spec. nov. — right gonopod of the holotype &, medial
view. (The solenomerite has moved out of its natural position). Fig. 12. K. decolorata spec.
nov. — right gonopod of the holotype &, medial view
216 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Lateral keels. — As in ocellata.
Sternites and legs. — Sternites and sternal process of the 5th segment as in
ocellata. Brushes of tibiae and tarsi as in hertitarsus and ocellata.
Anal segment. — Tail as in ocellata.
Gonopods. — (fig. 11). Femur and coxa of subequal length. Femoral process
arising from the basal part of the solenomerite proper, spiniform. Tibiotarsus
curving medio-distad, caudad and, finally, cephalo-proximad and somewhat laterad.
Female. — Unknown.
Remarks. — A similar remark as was made regarding the relationship between
ruficeps and hirtitarsus applies to ocellata and albonigra. These two species also
are very similar in colour and general morphology, although the gonopods are
differing importantly. In this respect it is interesting to note the similarity of the
gonopods of albonigra and ruficeps, two species otherwise rather disjunct.
A form closely related to albonigra was tepresented in the collection by a
single male specimen from East Borneo (Coll. Dr. E. MJOBERG, Mus. Stockholm).
It has a width of 2.3 mm, thus being considerably smaller than albonigra, and
differs furthermore in the gonopods. In these the femur is relatively smaller than
in albonigra and the tibiotarsus is making a wider curve. The rather poor state of
preservation of the animal and the absence of an exact locality renders a description
of this form useless.
Kalimantanina decolorata spec. nov.
Material. — West Borneo: Bungan River, July, 1894 (Borneo Exped., Coll.
Dr. A. W. NIEUWENHUIS, Mus. Leiden), 1 4 (holotype). East Borneo: Mahak-
kam River, Blu-u, November, 1898 (Borneo Exped., Coll. Dr. A. W. NIEUWEN-
HUIS, Mus. Leiden), 1 4.
Differing from ruficeps in the following characters.
Colour. — Possibly faded or not fully developed. Entirely pale brownish yel-
low. In the holotype only the 6th and a part of the 7th joints of the antennae
dark brown. The paratype has the proximal joints also brown.
Width. — Holotype 2.9 mm, paratype 3.0 mm.
Head and antennae. — Vertigial sulcus running downward to just below the
upper level of the antennal sockets.
Collum. — As in albonigra.
Body segments. — Pleural keels as in ocellata.
Lateral keels. — More prominent than in rwficeps and thus more resembling
those of ocellata except that the lateral margins when seen from the dorsal side
are somewhat less rounded. On the other hand the posterior edges are more
pronounced and rather narrowly rounded; in the keels of the 18th and 19th seg-
ments the posterior edges are minutely angular and projecting a little behind the
posterior margin of the segments.
Sternites and legs. — Sternites as in ocellata; the sternal process of the 5th
segment as in ##ficeps but more distinctly bilobate at the distal end. Brushes of
tibiae and tarsi as in ocellata.
Anal segment. — Sides of tail somewhat more concave at the base, scarcely
converging in the distal part. The end more distinctly emarginate than in ocellata.
C. A. W. JEEKEL : Paradoxosomatidae from Borneo Di
Gonopods. — (fig. 12). Femur about one and a half times as long as coxa.
Femoral process elongate-laminate, slightly constricted in the middle, arising about
halfway the femur and thus distinctly proximad of the base of the solenomerite.
Tibiotarsus comparatively small, curving medio-distad, caudad, proximad, and,
finally, cephalad and somewhat laterad.
Female. — Unknown.
Remarks. — Future collecting may show whether the specimen from Blu-u,
which differs from the type specimen only by the colour of the antennae, belongs
to a distinct subspecies.
Borneonina gen. nov.
Generic diagnosis. — 20 segments. Poreformula normal. Head without particu-
lars; labrum tridentate. Antennae rather long.
Segments rather strongly constricted. Prosomites with fine cellular structure,
somewhat dull. Waist rather broad, strongly longitudinally ribbed in the dorsal
half, finely and rather widely striate in the ventral half dorsal of the sternal level.
Metatergites shining, hairless or with very few hairs, without sculpture. Trans-
verse furrow distinct but not deeply impressed, finely longitudinally striate, present
from the 4th segment onwards. Pleural keels present on a number of anterior
segments.
Lateral keels weakly developed, posteriorly rounded in nearly all segments,
projecting behind the posterior margin of the metasomite in the 2nd segment only.
Keels of 2nd segment well below the level of those of the next segment.
Sternites longer than wide, scarcely to dispersedly setiferous, with moderately
developed cross impressions. Sternite of 5th segment with a well developed,
distally laminate process between the anterior legs. Sternite of 6th segment
somewhat modified only in the posterior part. Sternite of 7th segment with on
each side a latero-anterior sublongitudinal ridge-like protuberance. Legs rather
long, without distinct tibial or tarsal brushes.
Gonopod-coxa of moderate size, straight-cylindrical, latero-distal surface with
a setiferous area. Prefemur rather short, almost in one line with the femur and
laterally well demarcated from that joint, the demarcation almost transverse on
the longitudinal axis of the femur. Femur straight, well developed, somewhat
longer than the coxa. Postfemur weakly demarcated. Spermal channel running
straight along the medio-anterior side of the femur towards the base of the
solenomerite. Solenomerite rather short, flagelliform, arising from the anterior side
of the distal end of the femur. From the medio-anterior side of the distal end of
the femur, mesad of the course of the spermal channel arises a rather long,
lanceolate process. Tibiotarsus arising from the posterior side of the distal end
of the femur, comparatively small, distinctly demarcated from the femur. Tibio-
tarsus consisting of a simple solenophore, straight with only the distal end curving
a little cephalad. Lamina medialis and lamina lateralis both present, the latter
more strongly developed than the former, sheathing the greater part of the
solenomerite.
Type-species. — Borneonina retrorsa spec. nov.
Range. — Borneo.
218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Number of species. — One.
Remarks. — The general morphology of this genus is in close agreement with
that of Kalimantanina. Borneonina is separated particularly because of the dif-
ferent shape of the tibiotarsus of the gonopods. In B. retrorsa the tibiotarsus is
notably smaller than in Kalimantanina, and it is curved in a direction opposite to
that found in the latter genus. There seems to exist no particular relationship
between Borneonina and any of the previously described species of Sundanina.
Borneonina retrorsa spec. nov.
Material. — West Borneo: Bungan River, July, 1894 (Borneo Exped., Coll.
Dr. A. W. NIEUWENHUIS, Mus. Leiden), 1 4.
Colour. — Probably faded. Pale brownish yellow, with on each side on keel-
level a continuous brownish longitudinal band of the width of the poriferous keels.
There is also an indication of the presence of a continuous middorsal brownish
band, apparently widest in the middle of the pro- and metasomites.
Width. — 2.1 mm.
Head and antennae. — Labrum narrowly and rather weakly emarginate. Clypeus
moderately convex, moderately impressed towards the labrum. Lateral sides
scarcely emarginate. Surface of head smooth and shining. Clypeus and frons
rather dispersedly setiferous, vertex with two setae. Antennal sockets separated by
slightly more than the diameter of a socket, or by three fifths of the length of the
2nd antennal joint. Postantennal groove shallow, the wall weakly prominent.
Vertex moderately convex, not demarcated from the frontal area. Sulcus rather
weakly impressed, reaching downward to the upper level of the antennal sockets.
Antennae not particularly slender. Length of joints decreasing very slightly from
the 2nd to the 6th, the 6th joint about three quarters of the length of the 2nd.
Joints of subequal width, the 6th a little thicker than the others. Antennal joints
moderately to rather densely setiferous.
Collum. — (fig. 13—14). Slightly narrower than the head, subreniform in
dorsal outline. Anterior border very weakly rounded in the middle, somewhat
more strongly so towards the sides. Posterior border widely and weakly emarginate,
somewhat convex laterally. Lateral sides rather widely rounded, caudally with a
weak indication of a blunt posterior edge. Surface smooth and shining, longitudin-
ally as well as transversely moderately convex; a few hairs present. Lateral and
latero-anterior border with a fine marginal rim, disappearing gradually towards
the middle.
Body segments. — Metatergites mostly hairless. Transverse furrow present from
the 4th to the 18th segment, but weakly impressed on the 4th. Furrow reaching to
almost one third from the dorsal demarcation of the lateral keels in most segments.
Sides somewhat rugulose and dispersedly granular up to the 4th segment, smooth
and aimost without wrinkles from the 5th onwards. Up to the 4th segment the
pleural keels are well developed ridges, which are posteriorly produced into an
angular lappet which projects weakly behind the posterior margin of the somites.
On the 5th and the 6th segments there is only a very weak indication of the
presence of pleural keels near the posterior margin of the somites.
Lateral keels. — (fig. 13—16). 2nd segment scarcely wider than the collum.
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 219
Fig. 13—18. Borneonina retrorsa spec. nov. — 13: left side of the head and the four
anterior segments of the holotype @, lateral view. 14: left side of the head and the three
anterior segments of the holotype 4, dorsal view. 15: left side of the 10th and 11th
segments of the holotype 4, dorsal view. 16: the same, lateral view. 17: leg of the 7th
segment of the holotype 4. 18: right gonopod of the holotype &, medial view. p: femoral
process; 1: lamina lateralis; m: lamina medialis
220 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
The keels anteriorly somewhat shouldered at the base, the anterior and posterior
edges rounded, the posterior edge projecting slightly behind the border of the
segment. Marginal rim rather weakly developed. 3rd and 4th segments of sub-
equal width, slightly narrower than the 2nd. The keels anteriorly widely rounded,
posteriorly somewhat more narrowly rounded. Ventrally they are demarcated only
in their posterior half. 5th segment distinctly wider than the 4th. Keels from the
5th segment onwards anteriorly and posteriorly widely rounded, especially in the
poreless segments. Poriferous keels somewhat more prominent. In the posterior
half of the body the posterior edges of the poriferous keels become slightly more
prominent and in the 18th and 19th segments they are minutely rectangular. Dorsal
furrow of keels narrow, not reaching the waist. Seen from the lateral side the
poreless keels have their dorsal demarcation slightly concave, the poriferous ones
are weakly convex. Poreless keels ventrally demarcated only near the posterior end,
poriferous keels in the posterior half. Pores lateral in a slight excavation, a little
more near the ventral demarcation of the keels.
Sternites and legs. — Sternites 11/, X as long as broad, mostly sparsely seti-
ferous. Cross impressions distinct but not deep. Process of the sternite of the 5th
segment with the sides parallel at the base, slightly convergent in the distal half.
The process directed cephalo-ventrad and projecting slightly before the anterior
border of the sternite. End of process rounded and deeply incised medially, with-
out a brush of short setae, but normally setiferous. Behind the process a rather
deep transverse furrow; between the posterior pair of legs a normal longitudinal
impression. Sternite of 6th segment with a wide cross impression, the longitudinal
impression caudally widening triangularly. Sternite of 8th segment without
particulars. Legs (fig. 17) rather long; weakly to, in tarsi, moderately setiferous.
Anterior legs somewhat more densely setiferous but none of them with brushes.
Eenethyofmoints mes Alk
Anal segment. — Tail rather broad at the base, dorso-ventrally rather thick.
The sides moderately converging, straight; before the end a well developed
lateral setiferous tubercle. End of tail straight-truncate. Dorsal side of tail without
a transverse depression, ventral side somewhat concave. Valves with narrow rims,
the setae on small tubercles. Scale trapezoidal; the setiferous tubercles low and
not projecting.
Gonopods. — (fig. 18). Femur about one and a half times as long as coxa.
Lamina lateralis with an irregular margin.
Female. — Unknown.
Dajakina gen. nov.
Generic diagnosis. — 20 segments. Poreformula normal. Head without part-
iculars. Antennae of moderate length to rather long.
Segments rather weakly to rather strongly constricted. Waist distinctly ribbed.
Metatergites shining, hairless, without sculpture. Transverse furrow present from
the collum or from the 5th segment onwards. Pleural keels weakly developed in
a few anterior segments.
Lateral keels weakly developed. Only those of the 2nd segment projecting —
slightly behind the border of the metasomite, and situated well below the level
of those of the 3rd segment.
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 221
Sternites longer than or as long as wide, moderately to rather weakly setiferous,
with moderately developed cross impressions. Sternite of 5th segment of &
without process. Legs of moderate length to rather long. At least a few of the
anterior legs of the & with tarsal brushes.
Gonopod-coxa rather long, straight-cylindrical, the latero-distal surface with a
setiferous area. Prefemur somewhat elongate, slightly oblique on the axis of the
femur and laterally well demarcated from that joint, the demarcation almost
transverse on the longitudinal axis of the femur. Femur almost straight, well
developed, somewhat shorter than the coxa. Postfemur not demarcated. Spermal
channel running along the medial side of the femur towards the base of the
solenomerite, the latter arising from the anterior side of the distal end of the
femur. Solenomerite of moderate length, flagelliform. Tibiotarsus arising from
the posterior side of the femoral end, moderately developed, distinctly demarcated
from the femur. Tibiotarsus consisting of a simple solenophore curving widely in
a posterior direction. Lamina medialis and lamina lateralis well developed,
sheathing the solenomerite for its greater part. Lamina medialis with a laminate
process just proximad of the terminal end of the tibiotarsus.
Type-species. — Dajakina oculata spec. nov.
Range. — Borneo.
Number of species. — Two.
Remarks. — The erection of this genus, which besides the type-species also
includes Orthomorpha rotundata Att., may be regarded as a first step towards a
subdivision of the species so far included in the genus Orthomorpha Bollm. into
natural categories. These species all agree in having a relatively simple gonopod
structure, in which, for instance, the tibiotarsus is represented by a solenophore
without or with very small secondary processes.
The simple structure of the gonopods and the fact that the gonopod characters
in this group obviously belong to the more stable, which are subject only to minor
changes in the course of speciation, have somewhat obscured the heterogeneity of
Orthomorpha. However, several generic categories may be recognised when non-
gonopod characters as well as the smaller details of the gonopods, such as the
course of the spermal channel, the development of the laminae medialis and
lateralis of the tibiotarsus, etc., are taken into consideration.
A more extensive discussion on the mutual affinities of the species of Ortho-
morpha is given in connection with the redescription of O. beaumontii (Le
Guillou).
The genus Dajakina may be distinguished by a combination of characters of
which must be mentioned: the weakly developed lateral keels, the weakly developed
pleural keels, the absence of a sternal process in the 5th segment of the male,
the lack of a sharply demarcated postfemur in the gonopods, the what may be
called “normal” development of the laminae medialis and lateralis of the tibio-
tarsus of the gonopods and the conformation of the tip of the tibiotarsus.
Little can be said of the relationship of the genus, apart from the general
remark that it belongs to the Orthomorpha-Pratinus complex. The peculiar
emarginations of the posterior borders of the segments occurring in the type-
species of the genus remind of a similar structure described for Sundanina sbini-
222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Fig. 19—23. Dajakina oculata spec. nov. — 19: antenna of the holotype &. 20: left side
of the head and the three anterior segments of the holotype &, lateral view. 21: the same,
dorsal view. 22: left side of the 10th and 11th segments of the holotype 3, lateral view.
23: the same, dorsal view
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 223
pleura Carl from Birma and Sundanina emarginata Jeekel from Tonkin. In
particular S. emarginata seems to come rather close to D. oculata, in the shape of
these emarginations as well as in characters of the gonopods. In fact, it may be
more closely related to D. oculata than to the type-species of the genus Sundanina
and may represent the nearest relative of D. oculata outside Borneo. On the other
hand it seems to have sufficient characteristic features not to be united in the same
genus with that species.
Key to the species. — Contrary to the condition in Kalimantanina or Arthrogo-
nopus, the species of Dajakina can hardly be separated on their gonopod characters.
Other features, on the other hand, such as the shape of the lateral keels and
various secondary sexual characters of the males provide ample possibilities for
specific distinction.
1. Lateral keels of poriferous segments represented by low, more or less eye-
shaped protuberances which are sharply demarcated on all sides. Transverse
furrow of metatergites present from the collum to the 18th segment. Posterior
borders of the 8th to the 18th segments of the ¢ with emarginations in the
lower half of the sides and behind the sternites. Legs of the ¢, except the
last two pairs, with tarsal brushes. Width of ¢ 4.0 mm
LE MOO een D. oculata spec. nov.
— Lateral keels of poriferous segments low but normally ridge-like, only dorsally
sharply demarcated by a furrow. Transverse furrow of metatergites present
from the 5th segment onwards. Posterior borders of segments not emarginate.
Tarsal brushes of 4 present only in a few anterior pairs of legs. Width of
SALO TINTO ALIA IEA I Ne O VE D. rotundata (Att.)
00000000000
Dajakina oculata spec. nov.
Material. — West Borneo: Sambas, October, 1893 (Borneo Exped., Coll. Dr.
HALLIER, Mus. Leiden), 1 &.
Colour. — Possibly either faded or not fully developed: entirely pale brownish
yellow. The distal part of the 6th and the proximal part of the 7th joints of the
antennae dark brown.
Width. — 4.0 mm.
Head and antennae. — Labrum narrowly and rather deeply emarginate, tri-
dentate. Clypeus rather weakly convex, moderately impressed towards the labrum.
Lateral sides scarcely convex. Surface of head moderately shining, somewhat
irregularly rugulose in the clypeal part. Clypeal and frontal part of headplate
moderately to rather dispersedly setiferous, vertex with two bristles. Antennal
sockets separated by three quarters of the diameter of one socket, or by about one
third of the length of the 2nd antennal joint. Postantennal groove shallow, the
wall in front of it moderately prominent. Vertex moderately convex, demarcated
from the frontal area by a slight depression. Sulcus moderately impressed, with
some transverse wrinkles, reaching downward to just below the upper level of the
antennal sockets. (The lower end of the sulcus is marked by two minute con-
tiguous tubercles. However, this may be an anomalous structure since the frontal
area of the specimen studied was injured). Antennae (fig. 19) rather long, though
not particularly slender. Length of the 2nd to the Sth joints subequal, the 6th
224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
joint notably longer. Width of the 2nd, 3rd and 4th joints subequal, the 5th joint
somewhat narrower, the 6th conspicuously ventrally inflated. Antennal joints
sparsely to moderately setiferous.
Collum. — (fig. 20—21). Narrower than the head, subreniform in dorsal out-
line. Anterior border weakly rounded in the middle, slightly more strongly so
towards the lateral sides, laterally very weakly emarginate. Posterior border widely
and very weakly emarginate in the middle, weakly convex towards the sides. ~
Lateral sides rather widely rounded. Surface shining and somewhat uneven, a
moderately impressed transverse furrow at one third from the posterior border.
Surface hairless, weakly longitudinally and moderately transversely convex.
Lateral and latero-anterior border with a fine marginal rim gradually disappearing
towards the middle.
Body segments. — Rather weakly constricted. The waist rather narrow dorsally,
somewhat widening laterally; dorsal half rather finely but distinctly longitudinally
ribbed, ventral half, above the level of the sternites, finely striate. Prosomites
Fig. 24—25. Dajakina oculata spec. nov. — 24: leg of the 7th segment of the holotype 4.
25: right gonopod of the holotype ¢, medial view; lamina lateralis; m: lamina medialis
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 225
with fine cellular structure, rather dull. Transverse furrow present on all segments
up to the 18th, rather sharply and rather deeply impressed and in most segments
almost reaching the dorsal delimitation of the lateral keels. Sides with fine and
moderately dense granulation in all segments. Pleural keels up to the áth segment
represented by weak, curved, somewhat granular ridges on a slight swelling. On
the 5th segment only a swelling is visible. From the 8th to the 17th segments, and
also, though weakly, in the 18th segment, the posterior border of the metasomites
in the ventral half of the sides as well as ventrally behind the sternites is rather
strongly emarginate. As a consequence the posterior margin of these segments at
the level of the stigmata appears rather strongly produced posteriorly, the produced
part being somewhat inflated.
Lateral keels. — (fig. 20—23). 2nd segment distinctly wider than the collum.
The keels anteriorly shouldered, their anterior edge rather widely, their posterior
edges narrowly rounded; the posterior edge projecting a little caudad of the border
of the segment. Marginal rim present only in the anterior half of the keels. 3rd
and 4th segments of subequal width, distinctly narrower than the 2nd. The keels
anteriorly somewhat more narrowly rounded than posteriorly. They are demarcated,
dorsally only, by a narrow furrow. 5th segment distinctly wider than the 4th.
Keels of poriferous segments represented by low eye-shaped protuberances, sharply
demarcated on all sides. In posterior segments these protuberances become more
elongate towards the anterior side. Pores lateral, in a slight excavation, almost in
the middle of the keels. None of the keels angular. Poreless keels somewhat less
prominent than the poriferous, represented by low, rather short longitudinal ridges,
dorsally and ventrally demarcated by furrows, which do not reach the waist. Behind
the keels some irregular wrinkles curving upwards.
Sternites and legs. — Sternites one and a half times longer than broad,
moderately setiferous. Cross impressions well developed, rather wide, not sharply
impressed. 5th segment without sternal process, but with a normal cross impression.
Sternites of the 6th, 7th and 8th segments without particulars, except that in the
6th segment the sternite is slightly more concave between the posterior legs.
Legs (fig. 24) of moderate length in the anterior part of the body, becoming
distinctly more elongate in the posterior part. All legs distinctly incrassate, rather
weakly to moderately setiferous. Dense ventral brushes are present on the distal
part of the tibiae and on the tarsi of most of the legs. Brushes, in particular those
of the tibiae, thinning out in the legs of the posterior part of the body, almost
absent in the legs of the 17th and completely absent in the legs of the 18th
segment. Brushes consisting of typically penicillate hairs. Length of the joints of
(HN ESS MESSE Os 45-251.
Anal segment. — Tail rather broad and thick and rather long. Sides not con-
verging, even a little concave, before the end a weak lateral tubercle. End truncate
and scarcely emarginate. No dorsal transverse depression at the base of the tail.
Ventral side of tail hardly concave. Valves with narrow but rather high rims, the
setae on small tubercles. Scale triangular, the posterior edge rounded, the sides
almost straight. Setiferous tubercles low, not projecting.
Gonopods. — (fig. 25). Femur at the anterior side near the base with some
notches. Tibiotarsus curving weakly in a posterior and somewhat medial direction.
Female. — Unknown.
226 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Remarks. — A species very closely related to oculata was represented by two
female specimens from East Borneo: Blu-u, Mahakkam River, November 1898
(Borneo Exped., Coll. Dr. A. W. NIEUWENHUIS, Mus. Leiden). These females
differ from oculata in particular by the shape of the lateral keels of the 2nd seg-
ment. In these the anterior edge is acutely angular, the lateral margin straight and
the posterior completely absent. The two specimens are notably smaller than
oculata, having a width of 3.3 mm and 3.1 mm, respectively. The antennal sockets
are separated by somewhat more than the diameter of one socket. The antennae
are of moderate length, the 6th joint has a quite normal shape. The transverse
furrow is absent in the metatergites of the three anterior segments. The posterior
borders of the segments lack the emarginations as described for oculata. The legs
are of moderate length and width, and the tail is somewhat shorter than in
oculata.
Considering the characteristic shape of the poriferous keels these female
specimens must be closely related to oculata. It seems likely therefore that the
peculiar structure of the 6th antennal joint, the emarginations of the posterior
borders of the metasomites and the incrassate legs described here for oculata are
secondary sexual characters of the male.
Dajakina rotundata (Att.)
1931 Orthomorpha (O.) rotundata Attems, Zoologica, Stuttg., vol. 30, fasc. 3/4, p. 116,
FRS?
1937 Orthomorpha (O.) rotundata, Attems, Tierreich, vol. 68, p. 64, fig. 78.
Distribution. — West Borneo: Lebang Kara, Nanga Serawei.
Remarks. — The gonopods of this species, which was not represented in the
collection under report, appear to be almost identical with those of oculata.
Aside from some minor details they seem to differ only in the somewhat
stronger curved tibiotarsus and the comparatively slightly shorter femur. In view
of these scarcely significant differences it is highly interesting to note the impor-
tant dissimilarities in the external morphology of the two species.
D. rotundata is a much smaller species than oculata, the width being 2.5 mm, as
against 4.0 mm. The antennae are said to be of moderate length and width; no
statement was made on the structure of the 6th joint, so that this joint probably
lacks the conspicuous inflation described for oculata. The lateral keels of the
poriferous segments were described as being small, short and dorsally demarcated
by a furrow, which is not conform with the peculiar structure in oculata. Further-
more, the transverse furrow of the metatergites is present only from the Sth to
the 18th segments in rotundata. The characteristic emarginations of the posterior
border of the metasomites are not mentioned in the description of rotundata.
Finally, contrary to the condition found in oculata, the legs of rotundata are stated
to be long and slender, with the tarsal brushes present only in the anterior legs.
In all, the differences between oculata and rotundata reveal to what remarkable
extent the non-gonopod characters may diverge evolutionally, without this diver-
gence being paralleled by an important structural alteration in the gonopods.
C. A. W. JEEKEL : Paradoxosomatidae from Borneo DD
Arthrogonopus gen. nov.
Generic diagnosis. — 20 segments. Poreformula normal. Head without par-
ticulars; labrum tridentate. Antennae rather long.
Segments moderately (2 ) to rather strongly ( & ) constricted. Prosomites with
fine cellular structure, somewhat dull. Waist of moderate width ( 4) to rather
narrow (9 ), strongly longitudinally ribbed in the dorsal half, finely and rather
widely striate in the ventral half dorsad of the sternal level. Metatergites shining,
hairless or with very few hairs, without sculpture. Transverse furrow distinct but
not deeply impressed, finely longitudinally striate, present from the Sth segment
onwards. Pleural keels well developed on a number of anterior segments.
Lateral keels rather weakly developed. In nearly all segments posteriorly rounded
or minutely angular in a number of segments of the posterior half of the body.
Only the keels of the 2nd segment projecting behind the posterior margin of the
segment; eventually also the keels of the 19th segment very slightly projecting
behind the margin. Keels of 2nd segment well below the level of those of the
next segment.
Sternite with moderately developed cross impressions, longer than wide (4)
or about as long as wide ( © ), moderately to rather weakly setiferous. Sternite of
5th segment of & with a well developed process between the anterior legs.
Sternite of 6th segment of & somewhat modified: a little excavated especially in
the posterior part. Sternite of 7th segment of 4 with on each side a weak callous
protuberance before the gonopod opening. Sternite of 8th segment of & without
particulars. Legs rather long; without tibial or tarsal brushes in the ¢.
Gonopods with coxa of moderate size to rather small, almost straight-cylindrical;
latero-anterior side of the distal end with a setiferous area. Prefemur rather short,
slightly oblique on the longitudinal axis of the femur and laterally well demarcated
from that joint, the demarcation almost transverse on the longitudinal axis of the
femur. Femur almost straight, well developed, somewhat longer than the coxa.
In the basal half the femur has two laminate crests on the medial side. Postfemur
laterally sharply demarcated. Spermal channel running along the medial side of
the femur. In the basal part it runs along the posterior laminate crest towards the
posterior side of the femur; more distally it runs towards the anterior side of the
femur. At the medio-posterior side of the distal end of the femur a small femoral
process or knob may be present. Solenomerite of moderate length, flagelliform,
arising from the medio-anterior side of the distal end of the postfemur. Tibio-
tarsus springing from the posterior side of the distal end of the postfemur,
distinctly demarcated from that joint. Tibiotarsus moderately to rather weakly
developed, consisting either of a simple semicircular solenophore without acces-
sory processes or of a solenophore which on the lateral side gives rise to a compa-
ratively well developed secondary process. Lamina medialis absent, lamina lateralis
well developed. Solenomerite not sheathed by, but closely applied to the medial
side of the tibiotarsus.
Type-species. — Arthrogonopus edentulus nov. spec.
Range. — Borneo.
Number of species. — Four.
Remarks. — Besides the three new species described in this paper, this new
228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
genus also includes the one previously known as Orthomorpha borneona Att.
These four species are even more disjunct from the group of typical species of
Orthomorpha than the two united under Dajakina. They are characterised especially
by the absence of a lamina medialis in the tibiotarsus of the gonopods, by the two
laminate crests of the basal part of the femur of the gonopods and by the weakly
developod lateral keels of the metasomites. In the structure of the tibiotarsus and
its relationship to the solenomerite as well as in the weakly developed lateral keels
the new genus approaches the Sumatran species of Sundantna and the species of
Opisthodolichopus Verh. like O. javanicus (Att.), but Arthrogonopus may be
distinguished from these by the sharply demarcated postfemur of the gonopods.
Key to the species. — The species of Arthrogonopus have a quite monotonous
external morphology. The characters of diagnostic value are found mainly in the
gonopods.
1. Colour of the last two pairs of legs not differing from that of the others.
Gonopods with a small process at the medio-posterior side of the distal end of
the femur pointing mesad. Tibiotarsus with a secondary process arising from
the“lateralt side wine tes 40 Ana a. Mean. Ess see. ehe A. bifidus spec. nov.
— Last two pairs of legs infuscate at least distad of the middle of the tibiae, the
others brown only in the distal half of the tarsi. Gonopods without a femoral
process, or with a very small knob at the medio-posterior side of the distal end
of the femur. Tibiotarsus without a secondary process ........................ 2
2. Tibiotarsus of gonopods comparatively small, its greatest length about equal to
tworthirds of the length fof thes femur ma A. edentulus spec. nov.
— Tibiotarsus of gonopods larger, its greatest length about equal to that of the
Femmes | MEAN RE A REEN 3
3. Posterior edges of the lateral keels from the 5th segment onwards rounded.
Terminal end of tibiotarsus of the gonopods relatively short, acutely angular
(DSP ION OR RT A. borneonus (Att.)
— Lateral keels of the segments of the posterior half of the body with minute
acuminate posterior edges. Terminal lappet of tibiotarsus of the gonopods
longemmtounded@inuprotilom en ee A. denticulatus spec. nov.
Arthrogonopus edentulus spec. nov.
Material. — East Borneo: Blu-u, Mahakkam River, November, 1898 (Borneo
Exped., Coll. Dr. A. W. NIEUWENHUIS, Mus. Leiden), 2 ¢ (one of them the
holotype), 1 9. East Borneo: Upper Mahakkam River, 1899 (Borneo Exped.,
Coll. Dr. A. W. NIEUWENHUIS, Mus. Leiden), 1 &. East Borneo: Upper Ma-
hakkam River (Borneo Exped., Coll. Dr. A. W. NIEUWENHUIS, Mus. Leiden),
2.9. East (?) Borneo: S. Bilis, May, 1932 (Coll. H. R. H. PRINCE LEOPOLD OF
BELGIUM, Mus. Brussels), 1 4. West Borneo: Mt. Tilung, eastern slope, March
1899 (Borneo Exped., Coll. J. BÜTTIKOFER, Mus. Leiden), 1 4. West Borneo:
Gng. Kenepai, Pondok, January, 1894 (Borneo Exped., Coll. M. Moret, Mus.
Perden) sland:
Colour. — Difficult to ascertain because of the generally rather poor state of
preservation. In the holotype the head is castaneous, with the antennae very dark
brown, almost black, only the distal part of the 7th and the 8th joint whitish.
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 229
Fig. 26—30. Arthrogonopus edentulus spec. nov. — 26: left side of the head and the
three anterior segments of the holotype 4, lateral view. 27: the same, subdorsal view. 28:
left side of the 10th and 11th segments of the holotype &, dorsal view. 29: the same,
lateral view. 30: sternal process of the Sth segment of the holotype 4, lateral view
230 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Collum of a similar castaneous colour as the head. Subsequent body segments also
castaneous, but soon growing darker and from the 5th onwards very dark brown to
black, with the keels, in particular those of the poriferous segments yellowish
brown to brownish yellow. Venter brown. Sternites yellowish brown to
brownish yellow. Legs brownish yellow to yellow, the distal half of the tarsi a little
darker brownish. The two last pairs of legs distad of the middle of the femur
very dark brown or black. Anal segment brown, the tail entirely brownish yellow.
Valves dark brown, scale pale brown. The other ¢ from Blu-u is much paler,
being brown all over, but this may be due to preservation. The last two pairs of
legs are yellowish, black only distad of the middle of the tibiae. The other
specimens studied are mostly discoloured, but as far as could be determined their
colour was similar to that of the holotype. The 4 from S. Bilis is completely pale
brownish yellow, and probably freshly moulted. The ¢ from Mt. Tilung seems
to be well preserved. It has the head dark castaneous, darkest in the frontal area.
The colour of the antennae is as described for the holotype. The collum and the
subsequent body segments, including the lateral keels, are black. The venter dark
brown to black, with the sternites more or less dark brown. The legs pale brown
to yellowish, with the tips of the tarsi not notably darker. The last two pairs black
distad of the proximal quarter of the femur. Anal segment black or very dark
brown, only the scale paler brown.
Width. — Holotype ¢ 3.1 mm; other ¢ in the order of the above enumeration
3:2, mm 3.1 mm) 3/0 8mm, 3.2 mm, 3.1 mms Or in SOM Aln ar
Head and antennae. — Labrum narrowly and moderately emarginate. Clypeus
rather weakly convex, moderately impressed towards the labrum. Lateral sides
weakly and widely convex, with a notch above the labrum. Surface of head shining,
with some irregular wrinkles. Headplate rather densely to moderately setiferous in
the clypeal and frontal regions, four setae on vertex. Antennal sockets separated by
slightly more than the diameter of one socket or by about half the length of the
2nd antennal joint. Postantennal groove shallow, the wall in front moderately
prominent. Vertex moderately convex, not or only very weakly demarcated from
the frontal region. Sulcus rather weakly impressed, running downward to somewhat
below the upper level of the sockets. Antennae slender. Length of joints: 3 > 4
= 2 = 5 > 6, the 6th joint about as long as three quarters to two thirds of the
3rd. The 2nd, 3rd and 6th joints of subequal width, slightly thicker than the 4th
and 5th. Antennal joints moderately to rather densely setiferous.
Collum. — (fig. 26—27). Distinctly narrower than the head, subsemicircular
in dorsal outline. Anterior border very widely convex in the middle, somewhat
more narrowly rounded towards the lateral sides, weakly emarginate or straight
above the rounding of the lateral sides. Posterior border very weakly and widely
emarginate in the middle, laterally slightly convex and very weakly emarginate or
straight immediately above the lateral rounding. Lateral sides widely rounded.
Surface smooth and shining, moderately convex, slightly more so towards the sides.
A few setae may be present. Lateral and latero-anterior borders with a fine
marginal rim, disappearing gradually towards the middle of the anterior border.
Body segments. — Metatergites with some sparse hairs, in particular in the
anterior and posterior segments. Transverse furrow present up to the 18th, or,
eventually, up to the 17th segment, in the majority of the segments reaching to
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 231
Fig. 31—32. Arthrogonopus edentulus spec. nov. — 31: leg of the 7th segment of the
holotype &. 32: right gonopod of the holotype &, medial view. (The solenomerite has
moved somewhat out of its natural position)
about halfway the dorsal delimitation of the lateral keels. Sides dispersedly
granulate up to the 4th segment, smooth or very weakly wrinkled from the Sth
onwards. Pleural keels present up to the 7th segment, obsolete from the 8th
onwards. Up to the 4th segment they are represented by complete ridges which
are produced posteriorly into a well developed angular lappet projecting behinc
the posterior margin of the somite. In the 5th, 6th and 7th segments the pleura.
keels are represented by a small triangular lappet near the posterior margin of the
segments, which does not project behind that margin.
23 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Lateral keels. — (fig. 26—29). 2nd segment distinctly wider than the collum.
The keels anteriorly somewhat shouldered at the base, the anterior and posterior
edges rounded, the posterior edge projecting slightly behind the border of the
segment. Marginal rim present only along the anterior margin, laterally obsolete.
3rd and 4th segments of subequal width, somewhat narrower than the 2nd. The
keels anteriorly and posteriorly widely rounded, ventrally scarcely demarcated. 5th
segment distinctly wider than the 4th. Keels from the 5th segment onwards
anteriorly widely rounded, the posterior edges rather narrowly rounded, in par-
ticular in the keels of the segments of the posterior half of the body, but in none
of these angular. Poriferous keels somewhat more prominent than the poreless.
Dorsal furrow of keels narrow, not reaching the waist. Seen from the lateral side
the poreless keels have their dorsal margins slightly concave, the poriferous keels
are somewhat convex dorsally. Poreless keels ventrally demarcated near the
posterior end only, poriferous keels in the posterior half. Pores lateral, in a slight
excavation, almost in the middle between the dorsal and ventral demarcations of
the keels.
Sternites and legs. — Sternites one and a half times as long as broad, moderately
setiferous. Cross impressions rather sharp. Sternite of 5th segment (fig. 30) with
a thick process between the anterior legs, directed ventrad and somewhat cephalad
and projecting very slightly before the anterior border of the sternite. Lateral
sides of the process distinctly diverging in a distal direction. The end widely
rounded. Anterior side with a dense brush of short setae at the distal end. The
shape of the process appears to be somewhat variable. In some of the specimens
it differs from that of the holotype in that, from a lateral view, the distal end may
be somewhat more attenuate than illustrated. Sternite of Sth segment with a normal
transverse furrow and a normal longitudinal furrow between the posterior legs.
Sternite of 6th segment widely concave between the posterior legs, the coxae of
which are somewhat more widely separated than those of the anterior pair.
Transverse furrow almost obsolete, longitudinal furrow very wide. Legs (fig. 31)
moderately setiferous, but the ventral side, especially of the proximal joints, rather
densely setiferous. Last two pairs of legs distinctly longer than the preceding.
Length of joints73>6>5>4 >2>1.
Anal segment. — Tail of moderate length, rather thick at the base, of moderate
width, the sides proximally rather strongly, distally more weakly converging.
Before the end on each side a well developed setiferous tubercle. The end of
the tail truncate and weakly emarginate, the two terminal tubercles weakly
developed. Tail without distinct transverse depression at the base. Ventral side
weakly concave. Valves with rather narrow and rather low marginal rims. Seti-
ferous tubercles moderately developed. Scale subtrapezoidal or subtriangular, the
end slightly convex, the sides almost straight. Setiferous tubercles moderately
developed and not projecting.
Gonopods. — (fig. 32). Prefemur moderately developed. Femur without
femoral process or knob. Tibiotarsus comparatively small, its greatest length about
two thirds of the length of the femur. Tibiotarsus simple, without secondary
process, the end acuminate.
Female. — The 9 from Blu-u, although rather strongly discoloured, appears to
agree in colour with the holotype. The two © from Upper Mahakkam River have
C. A. W. JEFKEL : Paradoxosomatidae from Borneo 233
the collum castaneous, infuscate in the middle and along the margins. The tail here
is darker than in the holotype. Morphologically the 9 differ particularly in being
more robust than the ¢. The body segments are somewhat less strongly con-
stricted and the waist is relatively a little narrower. The development of the
pleural keels of the Sth, 6th and 7th segments is somewhat weaker, so that a
triangular lappet is not distinct in these segments. The sternites of the middle
segments are as long as wide to 11/3 X as long as wide. The legs are slightly
more slender; the pubescence, however, is practically similar to that of the legs of
themas
Remarks. — Variation of the specimens studied appears to be confined to the
coloration and the shape of the sternal process of the 5th segment of the males.
The taxonomic value to be attributed to these variations cannot be determined at
present in view of the small amount of material studied and because of its rather
poor condition. The gonopods of the males from different localities do not present
appreciable differences.
Arthrogonopus denticulatus spec. nov.
Material. — East Borneo: Blu-u, 30 September (Mus. Leiden), 1 & (holo-
type). East Borneo: Blu-u, Mahakkam River, November, 1898 (Borneo Exped.,
Coll. Dr. A. W. NIEUWENHUIS, Mus. Leiden), 4 ¢, 4:9. East Borneo: Long-
Blu-u, November, 1898 (Borneo Exped., Coll. Dr. A. W. NIEUWENHUIS, Mus.
Leiden), 1 &. East Borneo: Upper Mahakkam, 1899 (Borneo Exped., Coll. Dr.
A. W. NIEUWENHUIS, Mus. Leiden), 1 &. East Borneo: Upper Mahakkam River
(Borneo Exped., Coll. Dr. A. W. NIEUWENHUIS, Mus. Leiden), 1 &. East (?)
Borneo: June to August, 1900 (Borneo Exped., Coll. Dr. A. W. NIEUWENHUIS,
Mus. Leiden), 1 ¢. West Borneo: Gng. Kenepai, Pondok, January, 1894 (Bor-
neo Exped., Coll. M. Moret, Mus. Leiden), 1 2.
Differing from edentulus in the following characters.
Colour. — The holotype has essentially the same colour as the holotype of
edentulus, but the keels are black and the anal segment is castaneous. The other
4 have the head castaneous, the collum either completely castaneous or castaneous
with the margin and the middle infuscate. The subsequent segments are entirely
black, except the brown venter. The sternites and legs are brownish yellow, the
legs of the last two pairs black distad of the middle of the femur. Anal segment
entirely castaneous.
Width. — Holotype & 3.0 mm; other 4 in the order of the above enumeration:
mm 10m 3.0 mm; 3.0 mm, 3:0"mm, 3-2) mm, 3.4 mm, 3.0 mm: 2739
mm, 3.9 mm, 3.8 mm, 3.8 mm, 4.0 mm.
Body segments. — The transverse furrow of the metatergites extends slightly
further laterad than in edentulus. Pleural keels of the 5th to the 7th segments not
angular.
Lateral keels. — The posterior edges of the lateral keels are somewhat more
narrowly rounded. From about the 12th or 13th segment these edges become
angular and on subsequent segments (fig. 33) they are produced posteriorly in a
minute, sharp point. The posterior edges of the keels of the posterior segments
234
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
DA
Ad hy
"OG.
INR
/
hi
CARRI
Fig. 33—35. Arthrogonopus denticulatus spec. nov. — 33: left side of the 16th segment of
the holotype 4, dorsal view. 34: sternal process of the 5th segment of the holotype 4,
lateral view. 35: right gonopod of the holotype &, medial view. (The solonomerite has
slightly moved out of its natural position)
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 235
are not projecting behind the margins of the metasomites, but in the 19th seg-
ment they may attain the margin.
Sternites and legs. — Sternite of Sth segment (fig. 34) with the process
distally much more laminate than in edentulus. The lateral sides of the process
almost parallel or diverging only very slightly in the distal direction. Sternite of
6th segment more concave between the anterior legs than in edentulus, scarcely
raised above the ventral level of the metasomal ring between the posterior legs;
the transverse and longitudinal furrows are almost obsolete. Legs, especially those
of the anterior part of the body, somewhat more densely pubescent than in eden-
tulus.
Gonopods. — (fig. 35). Prefemur rather strongly developed. At the medio-
posterior side of the distal end of the femur a small knob. Tibiotarsus of moderate
size, the greatest length about equal to that of the femur.
Female. — The coloration of the females is more variable than that of the males.
Of four specimens from Blu-u, one has a colour similar to that of the males,
two others have pale spots in the middle of the metatergites, whilst the fourth has
the keels and the lateral sides pale, contrasting with the black dorsum. The 9
from Gng. Kenepai appears to be almost completely discoloured. Morphologically
the females differ from the males in a way very similar to that found in eden-
tulus. The lateral keels in the posterior segments are not pointed caudally as
strongly as in the males: really sharply pointed edges may be present only in a
few segments.
Remarks. — The colour variations of this species as represented in the material
under report are difficult to interpret. In part they seem to have been caused by
the generally rather poor state of preservation. However, the coloration of this
species and that of edentulus might be somewhat more variable than usually is
the case. Without the study of freshly preserved material it is impossible to give
an opinion on the systematic value of the variations.
The females of this species are very similar to those of edentulus. For the
distinction one has to rely in the first instance on the shape of the lateral keels
of the posterior segments. It is not possible to estimate the constancy of this
character since both species apparently were found to occur in the same localities.
It may be that some female specimens of denticulatus have been referred here to
edentulus on account of the absence of distinctly pointed posterior edges in the
lateral keels of the posterior segments.
Arthrogonopus borneonus (Att.)
1931 Orthomorpha (O.) borneona Attems, Zoologica, Stuttg., vol. 30, fasc. 3/4, p. 115,
fig. 174.
1937 Orthomorpha (O.) borneona, Attems, Tierreich, vol. 68, p. 67, fig. 84.
Distribution. — West Borneo: Mandai River, Mt. Rajah, Sungei Malang
Remarks. — This species, of which I did not see any material, comes very
close to denticulatus, in particular in the characters of the gonopods. However, the
distal end of the tibiotarsus of denticulatus appears to be more elongate and is
typically rounded, whereas in borneonus it is acuminate. The lateral keels in
236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
borneonus are stated to be rounded posteriorly in all segments, which is certainly
not the case in denticulatus. The relationship between borneonus and denticulatus
may be of a subspecific nature.
Arthrogonopus bifidus spec. nov.
Material. — West Borneo: Sambas, October, 1893 (Borneo Exped, Coll. Dr.
HALLIER, Mus. Leiden), 1 4.
Differing from edentulus in the following characters.
|
Fig. 36. Arthrogonopus bifidus spec. nov. — right gonopod of the holotype &, medial
view; p: femoral process; s: secondary process of the tibiotarsus
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 237
Colour. — Head dark castaneous, darkest in the frontal area. Antennae also
dark castaneous, with the tip of the 7th and the 8th joint white. Collum and body
segments very dark brown to black, the venter dark brownish grey. Sternites brown
to brownish yellow. Legs pale yellow to brownish yellow. Anal segment black,
the valves brown, the scale pale brown.
Width. — 2.9 mm.
Body segments. — The transverse furrow of the metatergites extending a little
more laterad than in edentulus. Pleural keels of the Sth to the 7th segments not
angular, similar to those of denticulatus.
Sternites and legs. — Sternal process of the 5th segment similar to that of
edentulus, but much more attenuate distally when seen from the lateral side. The
process directed a little more strongly cephalad, the distal end therefore projecting
somewhat more distinctly before the anterior border of the sternite. Sternite of
the 6th segment as in denticulatus.
Gonopods. — (fig. 36). Prefemur rather strongly developed. Femur with a
small femoral process at the medio-posterior side near the distal end, pointing
mesad. The process is small, elongate triangular, about one and a half times as
long as broad at its base. Tibiotarsus comparatively small, its greatest length about
equal to two thirds of the length of the femur. From the lateral side of the
tibiotarsus arises a laminate secondary process.
Female. — Unknown.
Remarks. — The close relationship between this species and the other species
of Arthrogonopus is clearly demonstrated by the great similarity in the general
morphology and particularly by the gonopod characters. A. bifidus, however,
seems to be somewhat disjunct by the presence of a small femoral process and a
secondary process in the tibiotarsus. The latter, which seems to have the function of
keeping in place the distal end of the solenomerite, may be a derivate structure.
Probably it is the homologue of the terminal end of the tibiotarsus in species like
denticulatus and edentulus, and the condition might be seen as the result of a
cleavage of the end of the tibiotarsus in the course of evolution.
Gigantomorpha gen. nov.
Generic diagnosis. — 20 segments. Poreformula normal. Head without part-
iculars; labrum tridentate. Antennae of moderate length.
Segments moderately (4) to rather weakly (2) constricted. Prosomites
shagreened or dulled by a very fine cellular structure. Waist narrow, distinctly
longitudinally ribbed or “beaded” in the dorsal half, smooth or finely striate
in the ventral half dorsad of the sternal level. Metatergites dull or more or less
shining, generally hairless, rugulose to subgranulose and with a transverse row of
more or less distinctly developed granules along the posterior margin and some
dispersed mostly very weakly developed granules in front of the transverse furrow.
Transverse furrow distinct but not deeply impressed, finely longitudinally striate,
present from the 4th or 5th segment onwards. Pleural keels present at least in
the segments of the anterior half of the body, moderately to well developed in
the anterior segments.
Lateral keels strongly to rather strongly developed, in all segments from the
238 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
2nd to the 19th with an angular, sometimes even spiniform posterior edge, which
often projects behind the posterior margin of the segments. Marginal rim of
keels narrow, in poriferous segments widening only in the area of the pore.
Keels generally on a high level, sometimes, especially in the males, projecting
somewhat above the middorsal surface of the metatergites. Keels of the 2nd seg:
ment either distinctly below the level of those of the next segment, or on the same
level.
Sternites with rather weakly developed cross-impressions, somewhat longer than
wide (3), as long as wide (3, 9) or somewhat wider than long (9),
moderately setiferous. Sternite of 5th segment of & with a moderately developed
process between the anterior legs. Sternite of 6th segment of 3 somewhat
modified: raised above the ventral surface of the metasomal ring between the
anterior legs only. Sternite of 7th segment of & with on each side before the
gonopod opening a rounded, somewhat granular process. Sternite of 8th segment of
& without modifications. Sternite from the 5th (2) or from the 8th (&) to
the 17th segment with a more or less developed pointed cone near the base of
each leg. Legs of moderate length to rather long. Brushes obsolete to very strong-
ly developed on all joints distad of the coxa, present only in the 4.
Gonopods with coxa of moderate size, almost straight-cylindrical; distal setiferous
area well developed. Prefemur somewhat elongate, slightly oblique on the long-
itudinal axis of the femur and laterally well demarcated from that joint, the
demarcation almost transverse on the femoral axis. Femur well developed, almost
straight, somewhat longer than the coxa, narrowing distad. In the basal half the
femur on its medial side with two more or less laminate crests. Postfemur laterally
sharply demarcated. Spermal channel running along the medial side of the femur. In
the proximal half it runs along the posterior laminate crest, or between the two
crests, towards the posterior side of the femur, turning in the distal half towards
the anterior side. Solenomerite of moderate length, flagelliform, arising from the
medio-anterior side of the distal end of the postfemur. Tibiotarsus springing from
the posterior side of the distal end of the postfemur, moderately developed,
distinctly demarcated from the postfemur. Tibiotarsus consisting of a simple
solenophore. Lamina lateralis well developed, lamina medialis developed only in
the distal part of the tibiotarsus, where it gives rise to a more or less complicated
laminate process. Tibiotarsus sheathing the solenomerite in its distal part only.
Type-species. — Gigantomorpha immanis spec. nov.
Range. — Borneo, Celebes.
Number of species. — Eight.
Remarks. — This genus, besides the new species described in the present paper,
includes two species previously referred to Pratinus Att., viz., Pratinus fasciatus
(Att.) from Borneo and Pratinus socialis (Carl) from Celebes. Like so many of
the species brought under the generic heading of Pratinus by ATTEMS in his
monograph of the ‘’Strongylosomidae’’ of 1937, these two only have a quite
remote relationship with the type of that genus. As a matter of fact, ATTEMS seems
to have been ignorant of the true identity of Pratinus, for a species closely
related to the type of that genus was described by him as Exphyodesmus greeni
Att. in 1936. In 1941, in following ATTEMS, also CHAMBERLIN referred a typical
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 239
species of Pratinus to the genus Euphyodesmus and even went as far as creating
a new subgenus for this species and greent. In short the synonymy of Pratinus is
as follows:
Pratinus Att.
1895 Prionopeltis Pocock, Ann. Mus. Genova, vol. 34, p. 828 (nom. praeocc.). Type-species:
P. cervinus Poc., by original designation.
1937 Pratinus Attems, Tierreich, vol. 68, p. 113. Type-species: P. planatus (Poc.), by
original designation.
1941 Euphyodesmus (Ceylonesmus) Chamberlin, Proc. Ent. Soc. Wash., vol. 43, p. 33.
Type-species: E. (C.) vector Chamb., by original designation.
Type-species. — P. cervinus (Poc.). There may be some controversy as to what
species should be regarded as the type of Pratinus. The name quite obviously has
been proposed as a substitute for Prionopeltis, judging from the indication “nom.
nov’. However, ATTEMS apparently overlooked the type designation by
Pocock, which, as a matter of fact, is somewhat concealed in the text of
Pocock’s paper, and selected planatus as the type species of Pratinus, thus taking
the first of the three original species of Prionopeltis. In view of the doubtless
intention of ATTEMS to substitute the name Prionopeltis rather than to erect a new
genus, it seems best to revert to cervinus as the type of Pratinus.
Range. — Ceylon, Burma, Tenasserim, Andaman Is.
Species. — Five: cervinus (Poc., 1895), greeni (Att., 1936), planatus (Poc.,
1895), taurinus (Poc., 1895), vector (Chamb., 1941).
Thus Pratinus is restricted to the original concept of Prionopeltis with the
addition of two misplaced species of Euphyodesmus. Its generic characters are to
be determined yet, although it seems clear that the genus may be regarded as inter-
mediate between Centrodesmus Poc. and Orthomorpha Bollm. From the latter
genus Pratinus is distinguished by the strongly developed, horn-like lateral keels,
the thin marginal rims of the keels and the remarkably condensed type of tibio-
tarsus of the gonopods. A sharply demarcated postfemur seems to be absent. The
differences between Pratinus and Centrodesmus, especially the Indochinese species
referred to that genus, are less obvious. The lateral keels in Centrodesmus are
even more strongly developed than in Pratinus, but it remains to be seen whether
or not this evolutional tendency is of sufficient importance to justify a generic
separation. Since, however, the Sumatran type-species of Centrodesmus was based
upon a female specimen and its more important characters are unknown, a dis-
cussion on the status of the genus and its boundaries becomes rather useless.
Aside from the true species of Pratinus enumerated above, the following have
been referred to the same genus in the “Tierreich”: P. fasciatus (Att.), P. socta-
lis (Carl), P. flaviventer (Att.), P. tenuipes (Att.), P. levigatus Att, P. rotundi-
collis Att., P. tuberculatus Att, P. glandulosus Att., P. arboricola Att. and P.
butteli (Carl). Besides, the following species were enlisted as “unsichere Arten”:
P. montanus (Chamb.), P. bicolor (Carl), P. haast: (Humb. & Sauss.), P. paviei
(Brol.), P. beaumontii (Att.)?, P. clarus (Chamb.) and P. dasys (Chamb.). All
of these species, except the six mentioned hereafter, will be discussed under
Orthomorpha.
As has been stated already, P. fasciatus (Att.) and P. socialis (Carl), for reasons
to be given below, are referable to the new genus Gigantomorpha. P. bicolor
240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
(Carl) and P. haast: (Humb. & Sauss.), both from New Zealand and both based
upon female specimens, are certainly not referable to any of the genera treated in
this paper. In fact, one may wonder whether their location in the family Paradoxo-
somatidae was correct. The presence of polygonal areas on the metatergites in the
two species points in the direction of the Sphaerotrichopodidae, a family, in contra-
distinction to the Paradoxosomatidae, rather well represented in the New Zealand
fauna. My attempt to verify this question failed, inasmuch as the type specimens of
the two species could not be found in the museums of Bern and Geneva
(respectively Dr. W. KUENZI, 12.1V.1955, and Dr. H. Gisin, 27.V.1955, in litt.).
P. clarus (Chamb.) from the Fiji Is. and Samoa is doubtless to be regarded as a
synonym of Chondromorpha xanthotricha (Att.), a species now known to have an
almost world-wide range. P. dasys (Chamb.) from the Fiji Is. may be either a
species of Chondromorpha or Anoplodesmus, possibly introduced from elsewhere.
Rather than to Pratinus, Gigantomorpha appears to be related to the group of
species assigned in the present paper to Orthomorpha. The general gonopod-
structure in Gigantomorpha is in close agreement with that of the typical species
of Orthomorpha, such as O. weberi (Poc.), O. hydrobiologica Att, etc. Dif-
ferences, however, are found in the presence of two laminate femoral crests, the
somewhat different course of the spermal channel and the presence of a laminate
or spiniform process at the distal end of the lamina medialis of the tibiotarsus in
Gigantomorpha. Other characters for distinguishing this genus from Orthomorpha
probably will be found in the shape of the marginal thickenings of the lateral
keels of the metasomites, in the development of the pleural keels, in the sternites,
legs, etc., but unfortunately many species of Orthomorpha are not sufficiently well
described as regards these characters.
Attention may be drawn to the similarity in the structure of the femoral crests
of the gonopods in Gigantomorpha and Arthrogonopus. This resemblance seems to
indicate a certain degree of relationship between the two genera, but other charac-
ters, either of the gonopods or of the external morphology, show this relationship
to be quite remote and fully justify a generic separation.
Among the species of Gigantomorpha are to be found the largest Paradoxosoma-
tidae yet described. The fact that these giants remained undiscovered for such a
long time, reveals once again the enormous amount of taxonomic work to be done
on tropical millipedes.
Key to the species. — As in related genera, the gonopod characters in the genus
Gigantomorpha offer few possibilities for the distinction of species. Characters of
diagnostic value are found mainly in the shape of the lateral keels, the development
of the pleural keels, the coloration, and various secondary sexual features of the
males.
The present key pertains only to the species from Borneo. The single species
known from Celebes needs a re-examination as regards several important characters.
1. Colour of body segments in both pro- and metasomites brown, without yel-
lowish mid-dorsal spots or band. Lateral keels entirely yellow. Lateral sides of
collum rounded, without an acute latero-posterior angle. Lateral keels of 2nd
segment turned somewhat downwards, distinctly below the level of those of
the 3rd segment. Proximal joints of the legs of the male normally setiferous,
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 241
the femora straight. Basal and distal lateral setiferous tubercles of the tail
weakly developed, terminal tubercles well developed. Anal scale with the
setiferous tubercles equalling or surpassing the middle of the posterior margin 2
Colour different. Sides of collum with an angular latero-posterior edge. Lateral
keels of the 2nd segment turned somewhat upwards, nearly at the same level
as those of the 3rd segment. Proximal joints of the legs of the male, especially
in the anterior half of the body, ventrally more or less densely setiferous, the
femora distinctly arched. Either the basal or the distal lateral tubercles of the tail,
or both, well developed; terminal tubercles more or less well developed. Anal
scale with vestigial or weakly developed setiferous tubercles, which do not
Surpass ithemmuddlenor the) posterior marsine nr ne nn 3
Along the posterior margin of nearly all segments laterally and ventrally an
irregular row of setae. Metatergites with rather small tubercles, of which there
are eight to ten behind the transverse furrow. Pleural keels strongly developed,
present up to the 17th segment. Lateral keels in none of the segments projecting
dorsad of the middle of the metatergites. Width ( 4,:9 ) 5.0 to 5.4 mm … …
I RO ISO ensaeleee G. trichopleura spec. nov.
Posterior margin of segments without setae. Tubercles on metatergites larger,
generally six behind the transverse furrow. Pleural keels somewhat less strongly
developed, present up to the 14th or 15th segment. Lateral keels somewhat
more strongly developed, slightly raised above the level of the middle of the
metatergites in a few posterior segments of the male. Width (4, 9) 6.3 to
SEO NN RI e G. spinescens spec. nov.
Colour of body segments very dark brown to blackish, with the lateral keels
and a rather narrow continuous middorsal band pale brownish. Femur of
gonopods rather strongly constricted about halfway. Process of lamina medialis
of the tibiotarsus rather weakly developed, acuminate. Terminal end of tibio-
CALSUSEMINCIVADIFI e en Sn NT tes Pere eee G. fasciata (Att.)
Body segments without a continuous middorsal pale band. Femur of gonopods
without a distinct constriction. Process of lamina medialis rather strongly
developed, the end truncate. Terminal end of tibiotarsus a rounded or
somemmatethianoullatal OPS nd 4
Colour of body segments uniformly black, with only the marginal rim of the
lateral keels near the posterior edges somewhat paler brown. Legs of the male
with the proximal joints set with short curved setae, the femora somewhat
MRC G. aterrima spec. nov.
Colour of body segments dorsally either pale brownish gray, or dark purplish
brown with a large yellowish spot in the middle of the prosomites and yellow
posterior edges in the lateral keels. Legs of the male with long erect setae, the
FEM OAN OLERSErON DRA ECE ANNAE: 5
Lateral keels of male raised above the middorsal surface of the metasomites in
the 3rd to the 19th segments. Legs of male slender, the proximal joints not
strongly incrassate and their ventral pubescence not brush-like … … … … … ….
BIN RUE ER N EERENS A dui bic, As G. cornalata spec. nov.
Lateral keels of male raised above the middorsal surface of the metasomites in
the 17th to the 19th segments only. Legs of male rather strongly incrassate,
especially the proximal joints. The ventral side of all joints distad of the
242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
coxa’ pwithadensembrushelikempubescencel tap. AE 6
6. Colour of body segments dorsally dark purplish brown, a broad median spot
on the prosomites and the latero-posterior edges of the lateral keels yellow.
Pleural keels disappearing in the 10th or 11th segment ... G. immanis spec. nov.
— Colour of body segments dorsally entirely pale brownish gray. Pleural keels
disappearing in the 16th or 17th segment ............... G. pallida spec. nov.
Gigantomorpha immanis spec. nov.
Material. — East Borneo: Mt. Tibang, 1400 m (Coll. Dr. E. MJÖBERG, Mus.
Stockholm), 2 ¢ (one of them the holotype). East Borneo: Pajan River (Coll.
Dr. E. MJÔBERG, Mus. Stockholm), 1 4, 19. East Borneo: Blu-u, Mahakkam
River, November, 1898 (Borneo Exped., Coll. Dr. A. W. NIEUWENHUIS, Mus.
Leiden), 1 4,2 9, 1 juv. 9. East Borneo: Blu-u, 25 September (Mus. Leiden),
1 4. East Borneo: Upper Mahakkam River, 1899 (Borneo Exped., Coll. Dr.
A. W. NIEUWENHUIS, Mus. Leiden), 2 9. East Borneo: Upper Mahakkam River
(Borneo Exped., Coll. Dr. A. W. NIEUWENHUIS, Mus. Leiden), 1 4, 1 fragm. of
3,3 9. East Borneo: Mahakkam (Borneo Exped., Coll. Dr. A. W. NIEUWEN-
HUIS, Mus. Leiden), 1 3.
Colour. — Head dark purplish brown, the clypeal area and a spot at the
posterior border of each of the antennal sockets pale yellow. Antennae pale yellow
to brownish yellow, the 6th joint infuscate towards the end, the 7th joint dark
brown, its distal end and the 8th joint whitish. Collum dark purplish brown, the
lateral keels and sometimes a weakly defined narrow band along the posterior
margin pale yellow. Body segments of same dark groundcolour as the collum. A
large middorsal spot on the prosomites, the latero-posterior edges of the lateral
keels and sometimes a weakly defined narrow band along the posterior margin
of the metatergites pale yellow. Venter pale brownish, sternites and legs pale
yellow. Anal segment dark purplish brown, the tail yellow, the margins of the
valves and the scale pale brownish.
Width. — Holotype & 7.2 mm; other 3 in the order of the above enumeration
7.0 mm; 7.2 mm; 6.0 mm; 6.5 mm; 6.6 mm; 6.6 mm; 6.6 mm; ı9 7.3 mm;
7.0 mm; 7.2 mm; 6.8 mm; 7.3 mm; 6.7 mm; 6.9 mm; 7.2 mm; juv. 9 with
19 segments 4.6 mm.
Head and antennae. — Labrum moderately emarginate. Clypeus rather weakly
convex, moderately impressed towards the labrum. Lateral sides widely and weakly
convex, somewhat emarginate or straight near the labrum. Surface of headplate
shining, slightly rugulose. Head moderately to rather dispersedly setiferous in the
clypeal and frontal region, vertex with four, two, or without setae. Antennal
Fig. 37—43. Gigantomorpha immanis spec. nov. — 37: left side of the head and the four
anterior segments of the holotype &, lateral view. 38: left side of the head and the three
anterior segments of the holotype 4, dorsal view. 39: left side of the 10th and 11th
segments of the holotype 4, dorsal view. 40: the same, lateral view. 41: left side of the
17th, 18th and 19th segments of the holotype 4, dorsal view. 42: leg of the 7th segment
of the holotype 4. 43: anal segment of the holotype &, ventral view
C. A. W. JEEKEL : Paradoxosomatidae from Borneo
243
244 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
sockets separated by 11/3 X the diameter of one of the sockets, or by about three
quarters of the length of the 2nd antennal joint. Postantennal groove shallow, the
wall in front moderately prominent. Vertex moderately convex, not or weakly
demarcated from the frontal area. Sulcus moderately impressed, running downward
to the upper level of the antennal sockets or just below that level. Sulcus with
some fine transverse wrinkles. Antennae not particularly slender. Length of joints:
3 > 2= 4= 5 = 6, or the 6th sligthly shorter and about three quarters of the
length of the 2nd. Joints of subequal width, only the 6th slightly thicker than the
others. Pubescence of antennae moderate to, distally, rather dense.
Collum. — (fig. 37—38). Somewhat wider than the head, subtrapezoidal in
dorsal outline. Anterior border very widely convex in the middle, somewhat more
narrowly rounded towards the sides, lateral border almost straight. Posterior border
scarcely emarginate in the middle, weakly convex more laterally; posterior border
of lateral keels somewhat emarginate. Latero-posterior edge acuminate, almost
rectangular. Surface shining, leathery rugulose, hairless, rather weakly convex. In
the middle at about one third from the anterior border a weak transverse depres-
sion. Along the posterior margin some low, flat, large tubercles are indicated,
generally represented by some rather coarse wrinkles. Lateral keels somewhat raised
to a horizontal level. Latero-anterior border with a marginal rim ending posteriorly
at some distance from the posterior border, gradually disappearing anteriorly
towards the middle of the anterior border.
Body segments. — Metatergites leathery rugulose, shining, hairless. Along the
posterior margin a row of generally six flat larger tubercles, which generally are
weakly developed and often resemble coarse wrinkles. Tubercles in the anterior
part of the metatergites vestigial. Transverse furrow present from the 5th to the
17th segments, generally reaching laterad to the base of the lateral keels and bifur-
cating there into two furrows or depressions, curving cephalad and caudad
respectively. Sides rugulose, rather densely and minutely granular up to the 4th
segment, more dispersedly granular from the 5th segment onwards. Pleural keels
present up to the 9th or 10th segment, obsolete from the 10th or 11th. Up to
the 4th segment they are represented by well developed, complete, granulate ridges
which are posteriorly produced into a conical process projecting a little behind
the posterior margin of the somites. From the 5th segment onwards the pleural
keels are represented by a well developed swelling which is rather densely
granulate above the anterior legs and a well developed pointed cone near the
posterior margin of the segment, directed caudo-laterad and not projecting behind
the margin. Both, swellings and cones, gradually diminish in size in subsequent
segments.
Lateral keels. — (fig. 37—41). 2nd segment somewhat wider than the collum.
The keels anteriorly somewhat shouldered at the base, the anterior margin rounded,
the lateral margin a little emarginate in the middle, the posterior edge acutely
angular and projecting rather strongly behind the posterior margin of the segment,
the posterior margin straight. Keels of 2nd segment directed somewhat upwards,
their level scarcely below that of the keels of the 3rd segment. Marginal rim well
developed along the anterior and latero-anterior borders; a very fine rim along the
posterior burder. 3rd and 4th segments of subequal width, somewhat narrower
than the 2nd. The keels anteriorly very weakly shouldered at the base, rather
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 245
widely rounded. The posterior edges projecting rather strongly behind the margins
of the segments. Keels directed upwards a little, those of the 4th segment less than
those of the 3rd. Premarginal furrow not reaching the posterior margin. Posterior
margin with a fine rim. Marginal rim ventrally not marked off by a furrow.
5th segment distinctly wider than the 4th. Keels from the 5th segment onwards
anteriorly and laterally widely rounded, scarcely or not shouldered at the base.
Posterior edges acutely angular, especially in the posterior segments projecting well
behind the margins of the segments. In the segments of the middle part of the
body, however, they hardly project behind the posterior margins. Keels horizontal
in most of the segments, only in the 17th or 18th to 19th segments they are raised
a little above the middorsal surface of the metasomites. Poriferous keels extending
very slightly more laterad than the poreless. Dorsal premarginal furrow reaching
from the waist to near the posterior border; a fine marginal rim along the posterior
border. Marginal rim ventrally generally demarcated only in the posterior half of
the poriferous keels. The rim generally not ventrally demarcated in the poreless
keels. Pores dorso-lateral in a slight excavation of the marginal rim. In most seg-
ments the pores are visible from the dorsal side.
Sternites and legs. — Sternites about as long as wide in middle segments. Cross
impressions represented by a rather sharp transverse furrow and a very weak and
wide longitudinal impression. Sternal process of Sth segment somewhat broader
than long, the sides almost parallel, the end truncate and scarcely incised in the
middle. In lateral view the process is thick, a little acuminate towards the rounded
end. Process directed ventrad, not projecting before the anterior border of the
sternite. Anterior side of process near the end densely set with short setae.
Sternite of 5th segment with a normal transverse furrow behind the process and
a normal longitudinal impression between the posterior legs; sternal cones at the
bases of the posterior legs vestigial. Sternite of 6th segment somewhat longitudinal-
ly convex and transversely concave between the anterior legs, without transverse
furrow and sternal cones. Granular protuberances of the sternite of the 7th segment
well developed. Sternites from the 8th to the 17th segments with rather well
developed pointed cones near the bases of the legs. The cones are pointing ventrad
and a little caudad. Legs (fig. 42) moderately setiferous, the ventral side of the
joints distad of the coxa with dense brushes up to the legs of the 17th segment.
The last two pairs without brushes, more normally setiferous. Legs rather strongly
incrassate, in particular the three proximal joints. From the 3rd or 4th pair onwards
the legs have the femora rather strongly arched. Legs of moderate length. Length
Ofsjoints: EU DETTE
Anal segment. — (fig. 43). Tail of moderate length, rather broad and thick at
the base. Sides rather strongly converging. Basal lateral setiferous tubercles rather
weakly developed, distal lateral setiferous tubercles and terminal tubercles modera-
tely developed. Near the base of the tail a more or less developed dorsal transverse
depression. Ventral side of tail somewhat concave. Valves with rather narrow rims,
the setiferous tubercles weakly developed. Scale parabolically rounded, the sides a
little concave, the setiferous tubercles rather weakly developed and not projecting
behind the margin.
Gonopods. — (fig. 44— 45). Coxal setiferous area extending from the lateral,
over the anterior towards the medial side. Anterior crest of the femur gradually
246 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Fig. 44—45. Gigantomorpha immanis spec. nov. — 44: right gonopod of the holotype 4,
medial view; 1: lamina lateralis; m: lamina medialis. 45: distal portion of the tibiotarsus
of the same; p: laminate process of the lamina medialis
fading away towards the distal end. Femur not constricted halfway. End of tibio-
tarsus with a small simply rounded lobe. Lamina medialis with a well developed
laminate process (p), which is T-shaped in cross section.
Female. — Aside from the usual secondary sexual characters the females differ
from males by the somewhat more robust build, caused by the weaker constriction
of the somites. Collum with a rectangular latero-posterior edge; the posterior
margins of the lateral keels of the collum scarcely emarginate. Dorsal surface of
metasomites a little more convex transversely, so that in none of the segments the
lateral keels project dorsad of the middle of the tergites. Lateral keels comparatively
a little less prominent, the posterior angles of the corresponding keels less acute
than in the males. Sternites somewhat longer than broad; sternal cones present
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 247
from the Sth to the 17th segments, somewhat more weakly developed than in
the males. Legs not notably shorter, but disinctly more slender, in particular in
the three proximal joints. Pubescence moderate, the ventral side of the joints
without dense brushes, but only slightly more densely setiferous than the dorsal
side. Femora straight. Length of joints: 3>6>5>2>4>1. The 6th joint
somewhat more distinctly longer than the Sth than in the males.
Juvenile. — The juvenile 9 with 19 segments is essentially similar to the adult
females, although most of the characters are less pronounced. In particular the
lateral keels are distinctly less prominent and generally do not project behind the
margins of the somites.
Remarks. — The specimens of zmmanis collected in the Mahakkam river area
differ in some minor details from the typical form from Mt. Tibang and Pajan
River. At least the males appear to have a slightly smaller size: 6.0 to 6.6 mm as
against 7.0 to 7.2 mm. Moreover the keels are very slightly less prominent because
their lateral margins are a little more parallel to the longitudinal axis of the body.
In the tail the basal lateral setiferous tubercles are decidedly more strongly develop-
ed. The setiferous tubercles of the anal scale are somewhat more strongly developed
too, so that they are projecting very slightly behind the posterior margin. However,
a much more abundant material is needed to decide whether these characters are
of really subspecific value or just represent population varieties.
A form not directly referable to zmmanis although very closely related to that
species was represented by a male and a female specimen from East Borneo,
without nearer location (Coll. Dr. E. MJ6BERG, Mus. Stockholm). Width of &
7.3 mm, of ® 7.6 mm. The male differs from that of zmmanis in the pubescence
of the legs. The dense brushes on the ventral side of the joints distad of the
coxa are thinning out in the legs of the second half of the body and disappear in
those of the 15th to the 18th segments. As regards the characters of the anal
segment both specimens agree with the zmmanis specimens from the Mahakkam
area. Without more material and an exact locality it seems best not to name this
particular form which may represent a subspecies of zmmanzis.
Gigantomorpha cornalata spec. nov.
Material. — East Borneo: Birang river (Coll. Dr. E. MjOBERG, Mus. Stock-
holm) 2e
Differing from immanis in the following characters.
Colour. — Entirely similar to that of zmmanis.
Width. — 5.8 mm in both ¢.
Head and antennae. — The 6th antennal joint scarcely shorter than the 2nd.
Collum. — (fig. 46—47). Width in relation to the head larger than in immanis.
Latero-posterior edge of keels somewhat more acute than in immanis, the posterior
border of the keels almost straight. Surface more finely rugulose to subgranulose.
Along the posterior border of the collum a few flat, rather indistinct tubercles
resembling coarse wrinkles. The keels slightly turned upwards.
Body segments. — Surface of metatergites more finely rugulose to subgranulose.
The tubercles along the posterior margin a little smaller than in immanis, but
slightly more prominent. Sides somewhat more densely and more coarsely granulate.
248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Fig. 46—51. Gigantomorpha cornalata spec. nov. — 46: left side of the head and the
four anterior segments of the holotype &, lateral view. 47: left side of the head and the
three anterior segments of the holotype 4, dorsal view. 48: left side of the 10th and 11th
segments of the holotype &, dorsal view. 49: left side of the 17th, 18th and 19th segments
of the holotype &, dorsal view. 50: leg of the 7th segment of the holotype &. 51: anal
segment of the holotype 4, ventral view
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 249
Pleural keels somewhat more strongly developed than in zmmanis. The anterior
swellings more coarsely granular, disappearing earlier than the posterior cones
which remain visible up to the 14th or 15th segment.
Lateral keels. — (fig. 47—49). More strongly developed than in smmanis. The
lateral margins more straight, diverging a little more strongly in caudal direction.
Posterior edges more acute, curving somewhat inwards in the posterior segments,
and projecting distinctly behind the posterior margin of the metasomites in all
segments. Keels of all segments slightly turned upwards, projecting dorsad of the
middle of the tergites from the 3rd to the 19th segments. Pores situated a little
more laterally, the excavation of the marginal rim around the pores more distinct.
Sternites and legs. — Sternal process of 5th segment slightly constricted at the
base. The distal end somewhat more rounded, though still weakly incised in the
middle. From a lateral view the process is more attenuate and the end more nar-
rowly rounded than in zmmanis. Legs (fig. 50) distinctly longer than in zmmanis,
the proximal joints not particularly incrassate. Ventral brushes considerably less
dense, the hairs longer. The brushes are gradually thinning out in the post-
gonopodial legs, but still remain rather dense even in the legs of the 17th segment.
Een sthwofsjoints 335-56 mdk
Anal segment. — (fig. 51). The tubercles of the tail are more or less equally
developed. Anal scale subtriangular, the posterior end more narrowly rounded.
Gonopods. — Similar to those of zmmanis. The anterior femoral crest more
abrubtly ending distally.
Female. — Unknown.
Gigantomorpha aterrima spec. nov.
Material. — East Borneo: Balikpapan, S. Mentawir, 1—11 October, 1950 (Coll.
A. M. R. WEGNER, Mus. Bogor), 3 4,1 2.
Differing from zmmanis in the following characters.
Colour. — Entirely black, with the clypeal area of the head, the posterior part
of the lateral margin of the keels, the venter and the end of the tail dark brown.
Antennae, legs and sternite pale yellowish to brownish yellow. The 6th antennal
joint infuscate towards the end, the 7th joint dark brown, but its distal part and
the 8th joint whitish.
Width. — Holotype 4 7.3 mm, other 4 7.5 and 7.8 mm;.9 7.8 mm.
Head and antennae. — 6th antennal joint of about the same length as the 2nd,
the 4th and the Sth; the 3rd somewhat longer.
Collum. — (fig. 52—53). Width in relation to the head larger than in zmmanzis.
Lateral sides of the keels very weakly emarginate in front of the latero-posterior
edge. Latero-posterior angle somewhat more acute than in immanis. Posterior
border of keels straight. Surface somewhat dull, much more finely rugulose than
in zmmanis. Tubercles at the posterior margin weakly indicated.
Body segments. — Metatergites somewhat dull, very finely rugulose. The tuber-
cles at the posterior margin distinct but rather small in most of the segments.
Pleural keels as in zmmanis, disappearing in the 10th or 11th segment, those of
the 4th segment not projecting behind the margin.
Lateral keels. — (fig. 52—55). Somewhat more strongly developed than in
250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
immanis. The posterior edges often sharply pointed and bent somewhat inwards,
the points especially distinct in the keels of the posterior segments. Posterior edges
in all segments projecting more or less behind the margin of the metasomites.
Keels either horizontal or turned a little upwards, in the latter case they project
dorsad of the middle of the metatergites either in a few anterior and posterior
segments or in nearly all segments. Marginal rim of the keels in all segments
distinctly narrower dorso-ventrally than in immanis, the dilatation around the pores
Fig. 52—56. Gigantomorpha aterrima spec. nov. — 52: left side of the head and the four
anterior segments of the holotype 4, lateral view. 53: left side of the head and the three
anterior segments of the holotype &, dorsal view. 54: left side of the 10th and 11th
segments of the holotype &, dorsal view. 55: left side of the 17th, 18th and 19th segments
of the holotype &, dorsal view. 56: leg of the 7th segment of the holotype &
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 251
smaller. Pores somewhat more lateral than in zmmanis, the excavation of the
marginal rim in the pore area more distinct.
Sternites and legs. — Sternal process of 5th segment similar to that of 777277241255,
but slightly narrower in lateral view. In posterior view the end is very
weakly incised, appearing like a wide V. Legs (fig. 56) similar to those of
immanis, very slightly less incrassate, the femora somewhat less strongly arched,
the ventral pubescence of the distal joints less dense. Ventral pubescence of the
proximal joints rather dense, at least in the legs of the anterior half of the body;
the setae are very short and typically curved distad. In the legs of the second half
of the body the pubescence is moderate, but the setae of the proximal joints remain
short. Length of joints: 3>6>5>4 >2>1.
Anal segment. — Tail almost as in cornalata. Anal scale as in zmmanis, the
setiferous tubercles a little more distinct.
Gonopods. — Similar to those of cornalata.
Female. — Aside from the usual secondary sexual characters the 9 differs
from the males in having a somewhat more robust appearance. Latero-posterior
edge of collum somewhat less acute than in the males. Dorsum of metatergites only
very little more convex. Lateral keels on a high level and in some anterior and
posterior segments almost projecting dorsad of the middle of the tergites. Keels
a little less prominent than in the males, the posterior edges scarcely less acute.
Sternites somewhat broader than long, the cones similarly well developed. Legs
slightly shorter but much more slender than in the males, moderately setiferous
and without a more dense ventral pubescence, the hairs normal. Femora straight.
enothvor joints > CESR ik
Gigantomorpha pallida spec. nov.
Material. — East Borneo: S. Mentawir, 13 October, 1950 (Coll. A. M. R.
WEGNER, Mus. Bogor), 3 &.
Differing from immanis in the following characters.
Colour. — Head rather dark brown, paler in the clypeal area. Antennae also
rather dark brown, the 6th joint infuscate towards the end, the 7th joint also dark,
but the distal end and the 8th joint whitish. Collum and the dorsal part of the
pro- and metasomites of the subsequent body segments pale brownish gray. In
the middle of the collum on each side straight behind the antennal sockets a
rather small rounded brown spot. In the body segments the lateral part of the
prosomites near the waist, the lateral part of the waist itself, the area of the meta-
somites before, immediately below and behind the lateral keels and the lateral
part of the posterior border of the metasomites dark brown. The middle portion
of the sides pale brownish gray. Venter, sternites and legs very pale brownish gray
or pale brownish. Anal segment pale brownish gray, the sides of the anal ring and
the valves dark brown.
Width. — Holotype 6.4 mm; other & 6.4 mm and 6.8 mm.
Head and antennae. — Length of antennal joints as in aterrima.
Collum. — (fig. 57). Width in relation to the head somewhat larger than in
immanis. Latero-posterior edges more acute, the posterior border of the keels
somewhat more distinctly emarginate. Surface somewhat dull, finely rugulose to
252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Fig. 57—60. Gigantomorpha pallida spec. nov. — 57: left side of the head and the three
anterior segments of the holotype 4, dorsal view. 58: left side of the 10th and 11th seg-
ments of the holotype &, dorsal view. 59: anal segment of the holotype &, ventral view.
60: femur of the right gonopod of the holotype 4, medial view.
sub-granulose, a few hairs may be present along the anterior border. Tubercles
weakly indicated.
Body segments. — Metatergites rugulose to sub-granulose. Tubercles along the
posterior margin small, but rather distinct. Tubercles in front of the transverse
furrow almost obsolete. Sides somewhat more coarsely and densely granulate.
Pleural keels more strongly developed than in zmmanis, the swellings more coarsely
granulate, posterior cones well developed, disappearing on the 16th or 17th
segment.
Lateral keels. — (fig. 57-58). Somewhat more strongly developed than in
immanis. The posterior edges often with a small sharp point curving a little in-
wards, in particular in the posterior segments. The edges slightly more acute,
scarcely or not projecting behind the margin of the segments in the middle part of
the body, but more strongly than in zmmanzs in the posterior segments. Pores more
lateral than in zmmanis, the lateral emargination of the rim in the pore area
somewhat more distinct.
Sternites and legs. — Sternal process of 5th segment distally acuminate from a
lateral view, the distal end sub-laminate. From a posterior view the base is very
slightly constricted, and the end is medially incised as in aterrima. Legs as in
immanis.
Anal segment. — (fig. 59). The basal and distal lateral setiferous tubercles of
nn un
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 253
the tail are distinctly more strongly developed. Setiferous tubercles of scale slightly
projecting.
Gonopods. — (fig. 60). The anterior femoral crest ends abruptly; the end
being even very slightly produced.
Female. — Unknown.
Gigantomorpha fasciata (Att.)
1898 Prionopeltis fasciatus Attems, Denks. Ak. Wien, vol. 67, p. 353, pl. 5, fig. 120—121
(i):
1912 Prionopeltis fasciatus, Carl, Rev. Suisse Zool., vol. 20, p. 141.
1937 Pratinus fasciatus, Attems, Tierreich, vol. 68, p. 115, fig. 150 (2).
Previous records. — Borneo (1); South Borneo: Bandjermasin (1), Pagat,
Barabei (2).
Material studied. — East Borneo: S. Mentawir, 1—13 October, 1950 (Coll. A.
M. R. WEGNER, Mus. Bogor), 4 &.
Differing from immanis in the following characters.
Colour. — Head black, brown in the clypeal area. Antennae very dark brown,
particularly in the distal joints. The tip of the 7th and the 8th joint whitish. Col-
lum and subsequent body segments black, with a rather narrow continuous pale
brown mid-dorsal band running from the anterior border of the collum to the end
of the tail. Lateral keels of collum and other segments with pale brown latero-
posterior edges. Venter brown. Sternites pale brown to brownish yellow. Legs pale
yellow. Anal segment. Sternites pale brown to brownish yellow. Legs pale yellow.
Anal segment laterally and dorsally black, aside from the mid-dorsal band. The
ventral side and the scale brown, the valves a little darker brown.
Width — 5:5: mm, 555mm 5.6. mm and! 5.6 mm.
Head and antennae. — Length of antennal joints as in aterrima.
Collum. — (fig. 61—62). Width in relation to the head only a little
larger than in immanis. Latero-posterior edges about rectangular, the posterior
border of the keels nearly straight. Surface somewhat dull, sub-granulose, a few
hairs along the anterior border. Tubercles obsolete. Lateral keels slightly raised
above the horizontal level.
Body segments. — Metatergites somewhat dull, sub-granulose. Tubercles along
the posterior margin small but distinct. Sides rather densely and somewhat more
coarsely granulate. Pleural keels more strongly developed, similar to those of
cornalata, disappearing in the 14th or the 15th segment.
Lateral keels. — (fig. 61-64). Very similar to those of immanis, but the
posterior edges somewhat more acute in the posterior segments. 2nd segment wider
than the collum, more distinctly than in zmmanis. All of the keels more or less
raised above the horizontal level, projecting above the middle of the metatergites
either in all segments from the áth to the 19th or only in a few anterior and
posterior segments. Pores distinctly more lateral, the marginal rim more distinctly
emarginate than in zmmanis.
Sternites and legs. — Sternites scarcely longer than wide. Process of the sternite
of the 5th segment distally acuminate from a lateral view, though not laminate.
From a posterior view the process is slightly constricted at the base, the end
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 255
scarcely emarginate. Legs (fig. 65) rather long and not particularly incrassate.
Ventral pubescence of the joints rather dense in the anterior legs, but gradually
thinning out, becoming moderately dense in the legs of the posterior half of the
body and normal in the last two pairs. Length of joints: 3> 6> 5> 4= 2>1.
Anal segment. — (fig. 66). Almost similar to that of zmmanis, but especially
the distal lateral setiferous tubercles of the tail somewhat more strongly developed.
Gonopods. — (fig. 67—68). The anterior femoral crest rather thick, less
laminate than in smmanis, ending more abruptly distally. Femur distinctly con-
stricted about halfway. Terminal lappet of tibiotarsus small. The end of the tibio-
tarsus curving laterad. The process of the lamina medialis rather weakly developed,
subspiniform.
Remarks. — It is not without some doubt that the presently described material
has been referred to fasciata, because the characters which actually have proved to
be of diagnostic value, such as the shape of the lateral keels, the rate of develop-
ment of the pleural keels, the shape and pubescence of the legs, etc., have scarcely
been mentioned by ATTEMS. Of the lateral keels of the type material of fasciata
only characters of generic value have been given, of the pleural keels we only know
that they are disappearing in the second half of the body, and of the legs it has
been said (vide CARL, 1912) that tarsal brushes are found only in the eight
anterior pairs of legs.
In the present material typical tarsal brushes are absent, although the tarsi of
the anterior legs are more densely setiferous, as usual indeed, than those of the
middle and posterior legs.
Still, there appears to be no evidence justifying a specific separation of the
present material. Obviously there is no important difference in coloration, the
type specimens apparently having been slightly paler. In outline, the gonopods of
the present specimens agree largely with the drawings published by ATTEMS,
although, there remain some minor discrepancies as a comparison of the figures
will show.
Although conforming essentially with the group of very closely related species
embracing immanis, cornalata, aterrima and pallida, fasciata is decidedly more
disjunct, in particular in the characters of its gonopods such as the femoral con-
striction, the weaker development of the process of the lamina medialis of the
tibiotarsus, etc.
Gigantomorpha trichopleura spec. nov.
Material. — East Borneo: without nearer location (Coll. Dr. E. MJOBERG,
Mus Stockholm) MEMOIRE 2ujuyaor
Differing from immanis in the following characters.
Fig. 61—68. Gigantomorpha fasciata (Att.). — 61: left side of the head and the four
anterior segments of a &, lateral view. 62: left side of the head and three anterior segments
of the same 4, dorsal view. 63: left side of the 10th and 11th segments of the same 4,
dorsal view. 64: left side of the 17th, 18th and 19th segments of the same 3, dorsal view.
65: leg of the 7th segment of the same &. 66: anal segment of the same 3, ventral view.
67: telopodite of the right gonopod of the same 4, medial view. (The solenomerite has
moved out of its natural position). 68: distal portion of the tibiotarsus of the same
256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Colour. — Head more or less dark castaneous, paler in the clypeal area. An-
tennae brownish yellow, the end of the 6th and particularly the 7th joint dark
brown. Tip of 7th and the 8th joint whitish. Collum and body segments more or
less dark castaneous, the lateral keels entirely yellow or brownish yellow. Venter
pale brown, the sternites and legs brownish yellow. Anal segment dark castaneous,
the tail yellowish, the valves and the scales paler castaneous.
Width. — Holotype & 5.0 mm; other 4 5.0 mm; 9 5.4 mm each juvenile 4
with 19 segments 3.1 mm, 3.1 mm and 3.2 mm; juvenile 9 with 19 segments
3.3 mm; juvenile 9 with 18 segments 2.5 mm.
Head and antennae. — Headplate rather weakly to sparsely setiferous in the
clypeal and frontal parts. Length of antennal joints as in aterrima.
Collum. — (fig. 69—70). Subsemicircular in dorsal outline. Sides rather widely
rounded, without a latero-posterior edge. Latero-anterior border almost straight
above the lateral rounding, posterior border with a weak notch above the lateral
rounding. Surface shining and rugulose as in smmanis, a few bristles along the
anterior margin, tubercles obsolete. The lateral margins slightly raised, the dorsal
surface moderately convex. Marginal rim rather weakly developed but extending
to the latero-posterior border.
Body segments. — Metatergites shining, rugulose, tubercles small but distinct,
generally 8 to 10 in front of the posterior border and 4 in front of the transverse
furrow. Tubercles sometimes setiferous. Transverse furrow indicated in the 3rd seg-
ment, generally well developed from the 4th to the 18th segments. Sides rather
weakly rugulose but rather coarsely granulate, in particular up to the 4th segment.
Along the posterior margin of the metasomites below the level of the keels an irre-
gular row of setae present up to about the 17th or 18th segment. Pleural keels present
up to the 17th segment, obsolete from the 18th onwards. They are well developed
and represented by complete, granulate to subtuberculate ridges up to the 6th
segment. From about the 7th segment the pleural keels consist of a well developed
longitudinal swelling which is densely granulate above the anterior legs and
granulate to subtuberculate above the posterior legs of each segment. A posterior
cone, directed laterad and somewhat caudad is well developed, but it does not
project behind the margin of the segments in the posterior half of the body.
Lateral keels. — (fig. 69—72). Anterior border of keels of 2nd segment
weakly rounded, latero-anterior edge with a small tooth. Lateral border widely
rounded, the margin somewhat irregular. Latero-posterior edge obtusely angular.
Marginal rim rather weakly developed, the pre-marginal furrow complete. Keels
of 2nd segment directed somewhat ventrad, their level distinctly below that of
the keels of the 3rd segment. Keels of the 3rd segment almost horizontal. From
the 4th segment onwards the keels are horizontal and do not project dorsad of the
metatergites. Pre-marginal furrow of the keels of the 3rd and 4th segments com-
Fig. 69—76. Gigantomorpha trichopleura spec. nov. — 69: left side of the head and the
four anterior segments of the holotype &, lateral view. 70: left side of the head and the
three anterior segments of the holotype &, dorsal view. 71: left side of the 10th and 11th
segments of the holotype 4, dorsal view. 72: the same, lateral view. 73: leg of the 7th
segment of the holotype 4. 74: anal segment of the holotype 4, ventral view. 75: right
gonopod of the holotype 4, medial view. 76: distal portion of the tibiotarsus of the same
C. A. W. JEEKEL : Paradoxosomatidae from Borneo
258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
plete. In all segments the lateral keels project behind the posterior margins of
the segments, especially in the posterior segments where the posterior edges of the
keels are more acute than in zmmanis and have a sharply pointed end curving in-
wards a little. Marginal rim of keels from the Sth segment onwards with a slight
lateral prominence about halfway followed up by an emargination. Pores lateral,
in most of the segments not visible from the dorsal side. Pore area rather weakly
emarginate.
Sternites and legs. — Sternites of middle segments about one and a quarter
longer than wide. Sternal process of the 5th segment distally acuminate when
seen from the lateral side, the end even sublaminate. From a posterior view the
process is not constricted at the base and the sides are parallel. The end has a very
wide V-shaped incision. Legs (fig. 73) moderately setiferous, more densely only in
the tarsi. Ventral side of the joints a little more densely setiferous than the dorsal
side but without brushes. Legs of moderate length, rather thick, the femora straight.
engthiot joints:53)> 6. — 2-4 eer.
Anal segment. — (fig. 74). Tail very broad at the base, the sides strongly con-
verging, a little concave. Basal and distal lateral setiferous tubercles very small,
terminal tubercles moderately developed. Dorsal transverse depression weak or
obsolete. Scale with well developed setiferous tubercles, projecting behind the
margin and equalling the middle of the posterior border.
Gonopods. — (fig. 75—76). The coxal setiferous area does not extend
to the medial side. Tibiotarsus comparatively smaller than in immanis, the terminal
end distinctly bifid. Process of lamina medialis less strongly developed, sub-
spiniform.
Female. — Aside from the usual secondary sexual characters the 9 differs
from the & in having a distinctly more robust appearance. The segments are
somewhat less constricted in the waist area and the lateral keels are distinctly less
prominent, their posterior edges somewhat less acute. Keels from the 3rd segment
to the segments of the middle of the body not projecting behind the posterior
margins of the segments. Sternites as long as wide. Legs a little more slender than
in the 4. Pubescence rather weak, that of the ventral side of the joints not dif-
ferent from that of the dorsal side. Length of joints: 6>3>2>4 = 5>1.
Juvenile. — The juvenile specimens agree essentially with the 9. The sculpture
of the collum and the metatergites is rather weakly rugulose, tubercles are missing,
but a few more hairs are present. The hairs along the lateral and ventral margins
of the somites are absent. The lateral keels are relatively much less developed than
in the adults.
Remarks. — Aithough trichopleura and the closely related spinescens are rather
disjunct from the other species of Gigantomorpha, there can be no doubt as to the
correctness of the present reference, since the gonopod structure in the two
species is basically the same as in immanis. In this respect, the genus G/gantomor-
pha is a perfect example to show the dubious value of the level of the lateral keels
of the 2nd segment as a taxonomic character. Doubtless previous authors merely
on account of this sole character would have referred zmmanis to the genus
Pratinus and trichopleura to Orthomorpha.
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 259
Gigantomorpha spinescens spec. nov.
Material. — East Borneo: without nearer location (Coll. Dr. E. MJÖBERG,
Mus. Stockholm), 2 4, 1 9.
Differing from zmmanis in the following characters.
Colour. — Exactly similar to that of trichopleura.
Width. — Holotype ¢ 6.8 mm; other ¢ 6.3 mm; 9 8.0 mm.
Head and antennae. — Pubescence of head as in trichoplenra. Length of anten-
nal joints as in aterrima.
Collum. — (fig. 77—78). Relative width of collum in comparison to the head
somewhat larger than in zmmanis. Dorsal outline subtrapezoidal to subsemicircular.
Fig. 77—81. Gigantomorpha spinescens spec. nov. — 77: left side of the head and the
four anterior segments of the holotype 4, lateral view. 78: left side of the head and the
three anterior segments of the holotype 4, dorsal view. 79: left side of the 10th and 11th
segments of the holotype 4, dorsal view. 80: left side of the 17th, 18th and 19th segments
of the holotype 4, dorsal view. 81: anal segment of the holotype 4, ventral view
260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Anterior border evenly rounded. Sides rather widely rounded, without a distinct
latero-posterior edge, although the lateral rounding is somewhat narrower at the
posterior side. Posterior border of the lateral sides straight, with a weak notch
above the lateral rounding. Surface somewhat dull, rather coarsely rugulose to
subgranulose. A few tubercles along the posterior border are either weakly in-
dicated or rather distinct. Lateral keels somewhat raised but not attaining a
horizontal level. Marginal rim laterally weakly developed, the furrow almost ob-
solete laterally.
Body segments. — Metatergites somewhat dull, rather coarsely rugulose to
subgranulose. Along the posterior margin six to, sometimes, eight rather large,
distinct tubercles. In the anterior segments there are also four large flat tubercles
behind the waist, which, however, rapidly grow less distinct on the subsequent
segments to become faintly indicated or obsolete in the segments of the middle
and posterior parts of the body. Transverse furrow of metatergites present, though
weakly developed, also in the 4th and the 18th segments. Sides rather coarsely
granular, in particular up to the 4th segment. Pleural keels more or less similarly
developed as in zmmanis but the anterior swelling somewhat more densely granu-
late; the posterior cones disappear in the 14th or 15th segment, the swellings
remain up to about the 17th segment.
Lateral keels. — (fig. 77—80). Keels of 2nd segment largely similar to those of
trichopleura. The latero-anterior edge without a tooth, but obtusely angular. Latero-
posterior edge about rectangular. Lateral keels of the other segments also largely
similar to those of trichopleura, the posterior edges more acute, becoming more
or less spiniform in the posterior segments. Keels mostly horizontal, only those
of the posterior segments raised slightly above the horizontal level but just not
projecting dorsad of the middle of the metatergites. Marginal rim of keels with a
similar lateral prominence, though much less distinct, than in trichopleura.
Sternites and legs. — Sternal process of the Sth segment differing from that of
trichopleura only in being somewhat constricted at the base. Legs as in tricho-
pleura, the ventral pubescence of the joints somewhat more dense.
Anal segment. — (fig. 81). Tail similar to that of trichopleura, the sides,
however, converging somewhat less strongly and not concave. Scale with setiferous
tubercles strongly developed, projecting well caudad of the middle of the posterior
margin.
Gonopods. — Similar to those of trichopleura.
Female. — Differing from the ¢ in the same characters as does the 9 of
trichopleura from the 3 of that species. Sternites somewhat broader than long.
Remarks. — A species very closely related to spinescens was represented by a
single female specimen from West Borneo: Sambas, October, 1893 (Borneo Exped.,
Coll. Dr. HALLIER, Mus. Leiden). It has a width of 7.2 mm and agrees with
spinescens in colour and general appearance. The main differences are found in
the smaller tubercles of the metatergites, the acute posterior edge of the lateral
keels of the 2nd segment, the much less elongate lateral keels of the posterior
segments and the development of the tubercles of the tail, which resemble those
of pallida.
In Celebes the genus Gigantomorpha is represented by a single species:
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 261
Gigantomorpha socialis (Carl)
1912 Prionopeltis socialis Carl, Rev. Suisse Zool., vol. 20, p. 139, pl. 5, fig. 13—15.
1937 Pratinus socialis, Attems, Tierreich, vol. 68, p. 116, fig. 151.
Distribution. — South Celebes: Bontorio.
In colour this species apparently resembles 77272475 and cornalata. In the char-
acters of the legs it seems to agree with zmmanis. The collum, however, has the
lateral sides rounded, the pleural keels remain visible up to the 16th or 17th
segment, so that soczalis is clearly distinct from both zmmanis and cornalata. The
gonopods of soczalis apparently have the generic characters of G/gantomorpha,
although the details of the acropodite have not been sufficiently illustrated.
Orthomorpha Bollm.
1859 Polydesmus (Paradesmus) De Saussure, Linn. Ent., vol. 13, p. 325.
1869 Polydesmus (Paradesmus), Humbert & De Saussure, Verh. zool. -bot. Ges. Wien, vol.
19, p. 670.
1893 Orthomorpha Bollman, Bull. U.S. Nat. Mus., vol. 46, P21595196:
1909 Orthomorpha, Pocock, Biol. Centr.-Amer., Diplop., p. 159.
1914 Orthomorpha (Orthomorpha), Attems, Arch. Naturg., vol. 80A, fasc. 4, p. 190.
1939 Asiomorpha Verhoeff, Zool. Anz., vol. 127, p. 117.
1944 Brasilogonopus Verhoeff, Arq. Mus. Nac. Rio de Janeiro, vol. 37, p. 274.
1958 Oxidus (Orthomorphina) Kraus, Senck. Biol., vol. 39, p. 100.
Type-species. — Polydesmus beaumontii Le Guillou, by subsequent designation:
Pocock, 1909. (Type-species of Astomorpha: Polydesmus coarctatus De Saussure,
by monotypy; of Brastlogonopus: B. attemsi Verhoeff, by monotypy; of Orthomor-
phina: Polydesmus coarctatus De Saussure, by original designation).
Range. — Burma, Indochina, Siam, Malayan Peninsula, Java, Borneo, Philip-
pines; one species has a circumtropical distribution.
Number of species. — 33 and one variety.
Remarks. — A discussion of the taxonomy of the genus has to be preceded by
some remarks relating the correctness of the application of the name Orthomorpha
and the designation of the type-species.
Orthomorpha was introduced by BOLLMAN in 1893 as a substitute for Parades-
mus Sauss., a name preoccupied by Paradesmus Corda. As such, Orthomorpha has
been generally accepted and used by all subsequent authors.
It is, therefore, rather surprising to find that the name Paradesmus Corda is
not included in the well-known Nomenclator Zoologicus by NEAVE. As a matter
of fact, in this Nomenclator we find behind the name Orthomorpha the addition:
“in err.?”, suggesting the probability of BOLLMAN’s name being superfluous.
Apparently, this condition recently has led to some confusion as regards the
correctness of the use of Orthomorpha and required some investigation.
Paradesmus has been used as a generic name first by A. J. C. CORDA in a
paper: “Ueber die Infusorien der Carlsbader Quellen”, published in: J. DE
Carro, Almanach de Carlsbad, 1835. Unfortunately, this paper was not available
to me. An abstract, however, was given by A. F. A. WIEGMANN in the “Bericht
über die Leistungen im Felde der Zoologie während des Jahres 1835” which
appeared in Arch. Naturg., vol. 2, fasc. 2, p. 184 (1836).
262 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Besides a discussion of CORDA’s paper by WIEGMANN, this article contains
also a synonymic list by C. G. EHRENBERG of the genera and species described
by Corba. In this list, Paradesmus Folium Corda is referred with some doubt to
the genus Fragilaria which belongs to the botanical Class Diatomeae. There seems
to be little doubt therefore that Paradesmus Corda was based on a vegetable
organism.
Contrary to this present day opinion, authors around 1835, or at least a number
of them, apparently were of the opinion that the Diatomeae (or Bacillaria, as they
were called at that time) were part of the animal kingdom. This is proved, for
instance, by the admission of Paradesmus Corda and other botanical genera, united
as a group Polygastrica, in the Nomenclator Zoologicus by AGassiz (1842—
1846).
Whether or not CORDA actually described Paradesmus as a zoological genus I
am unable to ascertain since I did not have access to his paper, but from WIEG-
MANN's discussion it may be inferred that CORDA regarded the organisms
described by him as animals. This question, however, becomes irrelevant, since
EHRENBERG ended his synonymic list, which included Paradesmus, with the
significant words: “Alle übrigen Formen kann ich nicht für Thiere anerkennen”.
From the available evidence I have concluded that Paradesmus Corda at least
for some time has figured as a generic name in zoology, so that according to ar-
ticle 2 of the International Code the substitution of Paradesmus Sauss. by
Orthomorpha Bollm. was quite correct.
There has been a considerable diversity of opinion as to the question which
species should be regarded as the type of the genus Orthomorpha.
In BOLLMAN’s paper of 1893 the name has been mentioned two times. The
author introduced it first on page 159, where in a footnote is stated: “Ortho-
morpha Bollman for Paradesmus Saussure, which is preoccupied”. On page 196
of the same paper the name is used again with the addition: “Orthomorpha, nom.
gen. nov. for Paradesmus Saussure, which is preoccupied by Paradesmus Corda,
Polyg.”. From this evidence one can only conclude that the name was proposed as
a direct substitute for Paradesmus Sauss. The type-species of Orthomorpha con-
sequently must be one of the species included in the original concept of Parades-
mus.
As far as I am aware a type-species of Paradesmus has not been designated
previous to 1893. Afterwards three opinions on this matter have been brought
forward.
The first of these has been the one of COOK and SILVESTRI, who in 1895 and
1896, respectively, without any comment, designated O. coarctata (Sauss.) as the
type of Orthomorpha, in which they were followed subsequently by ATTEMs, for
instance in the “Tierreich” in 1937.
The action by SILVESTRI was criticized by Pocock in 1909, with the following
cogent arguments:
“The name Paradesmus, Saussure, was originally applied to the following
species: — Section I. P. carolinensis; Section II. P. klugi, P. erchsoni, P. pictett;
Section IH. P. beaumonti (Linn. Ent, xiii, pp. 325, 326, 1859). One of these
must be its type, the addition of P. coarctatus to the list in 1860 (Mem. Soc.
Phys. Genève, xv, p. 297) not in any way affecting the question. As a matter
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 263
of fact this question was settled by Humbert and Saussure in 1869 (Verh. z.-b.
Ges. Wien, xix, p. 670), who eliminated the members of Sections I. and IT,
respectively under the names Ewryurus and Pachyurus, and restricted Paradesmus
to Section III., making P. beaumonti its type species*). But since Paradesmus was
preoccupied, Bollman proposed Orthomorpha to replace it. Therefore P. beaumonti
is the type of Orthomorpha and not P. coarctatus as Silvestri asserts (Ann. Mus.
Genova (2) xvi, p. 198, 1896).
Nothing needs to be added to Pocock’s words, which quite satisfactorily settle
the point.
In 1911, however, COOK rejected Pocock’s type designation, maintaining that
the Sections I and III of Paradesmus were considered by DE SAUSSURE as forming
transitions to other genera. According to Cook the name Orthomorpha could
therefore be applied only to the species of Section II. Fortunately Cook's
designation of P. klugii Brandt as the type of Paradesmus was antedated by that
of Pocock, for the acceptance of his proposal would have caused a lamentable
amount of confusion in nomenclature since the species of Section IT belong to the
family Euryuridae. As a matter of fact his opinion scarcely found any notice in
literature.
In the previous pages of this paper I have already emphasized the heterogeneity
which exists in the genera Orthomorpha and Pratinus as conceived by ATTEMS in
the Tierreich in 1937. From Orthomorpha two species, namely O. rotundata Att.
and O. borneona Att. have already been separated and brought to the new genera
Dajakina and Arthrogonopus, respectively. The genus Pratinus was brought back
to its original concept, and it was pointed out that most of the species which had
been referred to this genus by ATTEMS are true species of Orthomorpha. For
“Pratinus’ fasciatus (Att.) and “P.” socialis (Carl) and a number of newly
described species the genus Gigantomorpha has been erected.
The re-examination of the type-species of Orthomorpha gives a good opportunity
for a discussion of the taxonomic status of the remaining species of Orthomorpha
and Pratinus.
In 1937, ATTEMS recognized two subgenera of Orthomorpha: Orthomorpha
s. str. and Kalorthomorpha Att, 1914. Of course the latter name by isotypy is
an objective junior synonym of Oxidus Cook, 1911. In 1945, CHAMBERLIN has
already proposed to treat Oxidus as a separate genus, and in 1953, ATTEMS too,
although persistently rejecting CooK’s name, appears to have held the same
opinion as regards Kalorthomorpha. Although this proposal is quite correct in-
asmuch as the type-species of Oxidus, O. gracilis (C. Koch) seems to have no
obvious relationship with the type-species of Orthomorpha, it does not bring us
a step nearer to the solution of the main problem, the unraveling of the systematic
confusion existing in both genera.
As I have already stated in connection with the genus Gigantomorpha, many
of the species previously referred to the genus Pratinus are to be reallocated in
*) Here Pocock has been obviously mistaken, as HUMBERT and DE SAUSSURE did not
actually indicate a type-species; they merely referred beaumontii, together with coarctatus, to
Paradesmus in the restricted sense. Of course this does not affect the intention of his
argument.
264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
the genus Orthomorpha. Up to now the sole character used for the distinction of
Pratinus and Orthomorpha has been the position of the lateral keels of the second
segment. Species in which these keels are situated on about the same level as those
of the third segment have been referred to Pratinus, species in which these keels
are situated below the level of those of the third segment have been brought to
Orthomorpha. That this character is of comparatively little importance was shown
already in Gigantomorpha. Still, previous authors, namely ATTEMS and VERHOEFF,
have attributed to the character a primary systematic value even against the
evidence of the gonopods. The latter author even went so far as to base a family,
Prionopeltidae (= Pratinidae SCHUBART), on this single character. This family
quite correctly was discarded by ATTEMS as an artificial group in 1938 and 1953,
but curiously enough the latter author apparently would not risk uniting the
generic categories involved.
In 1953, ATTEMS erected the subgenus Paternostrana for the species of Ortho-
morpha in which the posterior edges of the lateral keels are rounded instead of
angular. The species referred by him to this subgenus were the following: O. ro-
tundata Att., constricta (Carl), hirtipes (Carl), borneona Att., sumbawana Att.,
javanica (Att.), thienemanni Att, and a new species minuscula Att. Although
ATTEMS was quite correct in separating these species from Orthomorpha, the
choice of the character upon which he based his subgenus was most unlucky, since
by evidence of the gonopod structure of the pertaining species it brings together
a number of quite unrelated forms.
As yet a type-species of Paternostrana has not been designated. To validate the
name, Orthomorpha sumbawana Att. is herewith selected as the type-species of
Paternostrana. Further reference to this genus will be found under category XI of
the discussion of the species of Orthomorpha.
I arranged the species of Orthomorpha s.str., Pratinus and Paternostrana sensu
ATTEMS, as far as they are not referred in the previous pages to other genera,
in a number of groups which are based in the first instance on the structure
of the gonopods.
Some of these groups may well prove to represent generic categories. For some
of them generic names are already available, and these have been used here,
although provisionally. For others generic names will have to be proposed; the
introduction of new names, however, is deliberately deferred to a later period,
pending actual study of the species involved.
The present arrangement is based largely upon literature data. It has been,
however, greatly facilitated by an inspection of the Pocock types in the British
Museum during a short visit there.
(1) — In this group are brought the species most closely associated with the
type-species of Orthomorpha, O. beaumontii (Le Guillou). A comparatively large
number of these hitherto have been referred to Pratinus, although others already
belonged to the genus according to the concept of previous authors. Some species,
previously considered of doubtful generic status, also are included, partly as a
result of the examination of the type material and partly on the evidence of
identified material.
In the species of this group the gonopods are of an extremely uniform type,
apparently changing very little during the course of speciation and failing to give
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 265
us reliable characters for specific distinction generally. The gonopods are
characterized by the laterally sharply demarcated postfemur and by a tibiotarsus in
which the lamina medialis and the lamina lateralis both are well developed,
sheathing the solenomerite for its greater part. The tibiotarsus lacks larger accessory
processes and terminates in a number of minute lappets, usually three. The
spermal channel runs along the medial side of the femur. However, characteristic
for the species of this group seems to be that in the prefemur and at the base of
the femur it follows a course along a typical fold in a more or less posterior
direction to bend abruptly distad when reaching the posterior side of the femur.
All the species of this group seem to have strongly developed lateral keels, with
the pores located laterally in a rather strongly thickened marginal rim.
The following species belong here.
ee acutangulus (Newport, 1844) Philippines
arboricola (Att, 1937) Indochina
— beaumontu (Le Guillou, 1841) Borneo
- bipunctata (Sinclair, 1901) Malacca
— cambodjana (Att., 1953) Indochina
= clivicola Poc., 1895 Burma
— coarctata (Sauss., 1860) Circumtropical
— coarctata var. gigas Att, 1927 Teun Id., Banda Sea
— consocius Chamb., 1945 Java
— conspicua (Poc., 1894) Java
— flaviventer (Att., 1898) Java
— francisca Att., 1930 Lombok
— fuscocollaris Poc., 1895 Tenasserim
— gestri Poc., 1895 Tenasserim
— glandulosa (Att, 1937) Indochina
— granosa (Att., 1953) Indochina
— hydrobiologica Att., 1930 Java, Indochina
== insularis Poc., 1895 Tenasserim
— intercedens Att., 1937 Burma, Tenasserim
-- karschii (Poc., 1889) Mergui, Tenasserim
— montana (Chamb., 1921) Philippines
— monticola Poc., 1895 Burma
— oatesti Poc., 1895 Tenasserim
— palonensis Poc., 1895 Burma
— paviei Brol., 1896 Siam
— rotundicollis (Att., 1937) Indochina
— spinala Att., 1932 Karimon Djawa
— tenuipes (Att, 1898) Java
-—- tuberculata (Att, 1937) Indochina
— unicolor (Att., 1930) Java
— weberi (Poc., 1894) Java
— zehnineri Carl, 1902 Java
The type specimens of O. acutangulus (Newp.) in the British Museum, a male
and a fragment of a female specimen in the dry collection, have been examined.
The species clearly belongs to the present group, although the tibiotarsus in both
266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
gonopods is broken off. It is one of the species which previously would have been
referred to the genus Pratinus, the keels of the 2nd segment being on a high level.
For the reasons why O. beaumontii belongs here, the reader is referred to the
remarks on page 272 relating to that species.
In the following two categories some species are brought together which although
probably rather closely related to those of the first group seem to be somewhat
disjunct by evidence of the gonopod structure. Their taxonomic position remains
somewhat uncertain.
(II) — The single species of this category differs from the species of group
(1) by having a comparatively shorter gonopod femur and by the apparent lack of
a sharply demarcated postfemur. The distal end of the tibiotarsus, however, seems
to be similar to that in the more typical species of Orthomorpha, and the species
may prove after re-examination to be only a somewhat aberrant form of the first
group.
Orthomorpha butteli (Carl, 1922) Java
(III) — This group also consists of a single species which, if we may trust
CARL's drawing of the gonopod, lacks a sharply defined postfemur. Moreover, the
distal end of the tibiotarsus of the gonopods differs from that of the species of
the first group by being more deeply split.
Orthomorpha bipulvillata Carl, 1902 Java
The following groups consist of species considered to be either subgenerically
or generically distinct from the species of group (1). For some of these categories
generic names are already available, for others new names will have to be proposed,
an action, however, which better be deferred until actual study of the species
involved.
(IV) — The single species of this group was referred to the genus Pagioprium
by ATTEMS in 1937. In the meantime, however, it was shown (JEEKEL, 1951)
that it is not congeneric with the type-species of that genus, a species now be-
longing to the genus Tectoporus Carl. In the gonopods “O.” cornuta differs from
the species of group (1) by the characteristically notched lamina medialis of the tibio-
tarsus, and apparently also by the absence of a sharply demarcated postfemur. The
smooth waist of the body segments, the absence of pleural keels and the typical
structure of the margins of the lateral keels also may prove to be characters of
subgeneric or generic value.
“Orthomorpha’ cornuta Att, 1930 Sumbawa (and Sabang?)
(V) — This category also consists of one species which apparently has no
distinct demarcation between the femur and the postfemur of the gonopods, and
has a characteristically reduced tibiotarsus. The lateral keels seem to resemble
those of the species in the first group, but the waist of the body segments has no
sculpture and pleural keels are missing.
“Orthomorpha’ anastasia Att., 1930 Lombok
(VI) — The two forms referred to this group present another instance of
the taxonomic confusion which has resulted from using the position of the
lateral keels of the 2nd segment as a discriminating character. The first form was
described as a Pratinus, the second as an Orthomorpha, but if it were not for some
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 267
small discrepancies in the descriptions, one would be inclined to unite both on
account of the practically complete identity of the gonopods.
Both species lack a demarcation between femur and postfemur in the gonopods,
and furthermore differ from the species of the first category in the structure of
the marginal rim of the lateral keels. The tibiotarsus has not been described clearly,
but seems to have a typical structure and way of sheathing the solenomerite.
“Orthomorpha’ exarata (Att, 1953) Indochina
— setosa Att., 1937 Indochina
(VII) — Referable to this category is a single species, which seems to have
sufficient characteristic features to justify a generic separation from Orthomorpha.
As a matter of fact, it was made the type-species of a genus Letozonius by ATTEMS
in 1953, after having been referred to Pratinus originally. Lezozonius, which, as
far as I am aware, was in fact a nomen nudum, is characterized in the gonopods
by the structure of the tibiotarsus in which the lamina medialis seems to be un-
usually strongly developed as against the somewhat reduced lamina lateralis, so
that the solenomerite appears to be applied to the lateral side of the tibiotarsus, a
condition somewhat reminiscent of what is found in the genus Tectoporus. Generic
characters also may be the absence of a distinct postfemur in the gonopods, the
smooth waist of the body segments and the absence of a transverse furrow on the
metatergites.
Leiozonius levigatus (Att., 1937) Indochina
(VIII) — In 1953 ATTEMS described a genus Piccola which was based on a
single species, P. odontopyga. Apparently on account of the position of the lateral
keels of the 2nd segment, the genus was compared with Pratinus and separated
from it by the absence of well developed lateral keels, the absence of a sternal
process in the 5th segment of the male, the irregularly wrinkled waist and the
long terminal tubercles of the tail. However, a comparison of the gonopod structure
of P. odontopyga with that of certain species referred to the genus Orthomorpha
shows once again that too much value has been given to the characters of the
lateral keels of the 2nd segment. Indeed, P. odontopyga is closely associated with
five species previously referred to Orthomorpha. The whole group is characterized
by the absence of a sharp demarcation between femur and postfemur of the
gonopods, by the weak development of the lamina medialis of the tibiotarsus, and,
apparently by the presence of a typical laminate crest on the medial side of the
basal half of the gonopod femur.
Piccola banana (Att., 1937) Indochina
— corrugata (Att., 1953) Indochina
— debilis (Att, 1953) Indochina
— minuscula (Att., 1953) Indochina
— odontopyga Att., 1953 Indochina
— spadix (Att., 1937) Indochina
(IX) — The three species of this category agree in having the lamina medialis
of the tibiotarsus very weakly developed to almost obsolete, whereas the lamina
lateralis is well developed. Consequently the solenomerite is more or less freely
applied to the medial side of the tibiotarsus, a condition strongly suggesting that
of the species of Arthrogonopus or the Sumatran species of Sundanina. The
solenomerite in its natural position has its end near a small more or less triangular
268 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
lobe of the tibiotarsus, just as for instance in Arthrogonopus edentulus or A.
denticulatus. A postfemur is not indicated. The known species have the lateral
keels rather weakly developed.
VERHOEFF proposed in 1941 the genus Opzsthodolichopus for one of the
species of this group, viz., thienemanni, a generic name which now is adopted
for the following species.
Opisthodolichopus javanicus (Att., 1903) Java
— nigricornis (Poc., 1894) Sumatra
— thienemanni (Att. 1930) Sumatra
(X) — A species, possibly related to the previous group but yet insufficiently
known, may be provisionally located here. The tibiotarsus of the gonopods appears
to have a somewhat different structure.
’Orthomorpha” semicarnea (Poc., 1894) Sumatra
(XI) — This category contains only two species, which are closely related if
not identical. The tibiotarsus of the gonopods has a strongly developed lamina
medialis, whereas the lamina lateralis appears to be almost completely absent.
The distal part of the solenomerite consequently is almost freely applied to the
lateral side of the tibiotarsus as in Leiozonius. There appears to be, however, no
close relationship between this group and Le/ozonius judging from the differences
in the general outline of the gonopods. A postfemur is not demarcated, the lateral
keels are rather weakly developed and both forms reportedly lack pleural keels.
To this group the genus Paternostrana Att., 1953, has been restricted. Possibly it
is related to Tectoporus Carl.
Paternostrana sumbawana (Att., 1930) Sumbawa
— vinosa (Poc., 1894) Flores
(XII) — The gonopod structure of the two species enumerated here is not
sufficiently known and a positive statement on their status is therefore premature.
However, the outline of the tibiotarsus seems to be quite distinct from that met
with in any of the other categories.
“Orthomorpha’ constricta (Carl, 1912) Celebes
-- hirtipes (Carl, 1912) Celebes
(XIII) — The species of this group certainly deserves a generic separation from
Orthomorpha. Actual study of specimens of the original series has shown that it
is probably related to species like Sundanina sigma Att., 1953, on the one hand and
Anoplodesmus mutilatus Att., 1953, on the other, although it lacks the processes
in the postfemoral region of the gonopods. The postfemur is demarcated from the
femur, but not as distinctly as in the more typical species of Orthomorpha. The
tibiotarsus is more elongate than in any of the species of the previous groups,
and has a normally developed lamina medialis and lamina lateralis.
“Orthomorpha’ doriae (Poc., 1895) Burma
(XIV) — A number of species referred to Orthomorpha by previous authors
are characterized by the presence of a rather large lanceolate or spine-like process
arising from the lamina medialis of the tibiotarsus of the gonopods, more or less
similar to what is found in the genus G/gantomorpha. These species, enumerated
under the present category, may not form an entirely homogeneous group, although
at least there appears to be some geographical coherence. Generally a postfemur
is rather weakly demarcated. Only in “O.” harpaga the demarcation appears to
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 269
be quite distinct, and also by other characters this species may be somewhat disjunct
from the others, having some affinities to the species of category (I).
A brief examination of specimens of the typical series of melanopleuris, min-
lana and miranda has shown a close relationship of these three forms with festiva,
orophila and uncinata. However, a thorough study of the species of this group is
needed to establish its characters. They certainly seem to be generically distinct
from the species of group (1).
For one of the species, miranda, a generic name was proposed by SILVESTRI as
far back as 1896. This name, Brachytropis, is preoccupied.
“Orthomorpha’ festiva Brol., 1896 Siam
— harpaga Att., 1937 Indochina
— mediovirgata Carl, 1941 Burma
— melanopleuris Poc., 1895 Burma
— minlana Poc., 1895 Burma
— miranda Poc., 1895 Burma
— orophila Carl, 1941 Burma
— uncinata Att., 1931 Siam
There seems to be good reason to believe that the following species, not
represented in the collection of the British Museum, also belong to this group.
“Orthomorpha’ bistriata Poc., 1895 Burma
— bivittata Poc., 1895 Burma
— comotti Poc., 1895 Burma
— pardalis Poc., 1895 Burma
(XV) — Aside from the process of the lamina medialis also present in the
previous group, the three following species have in addition a more basal spini-
form process on the medial side of the tibiotarsus. The postfemur appears to be
clearly demarcated. Of these species, coriacea has been referred to the subgenus
Kalorthomorpha by ATTEMS.
“Orthomorpha’ armata Carl, 1902 Java
—- atrorosea (Poc., 1894) Java
— coriacea Carl, 1902 Java
Orthomorpha beaumontii (Le Guillou)
1841 Polydesmus Beaumontii Le Guillou, Bull. Soc. Philom. Paris, 1841, p. 85.
1847 Polydesmus Beaumontii, Gervais, in: WALCKENAER & GERVAIS, Hist. Nat. Aptères,
vol. 4, p. 101.
1859 Polydesmus (Paradesmus) Beaumontii, De Saussure, Linn. Ent. vol. 13, p. 325.
1869 Polydesmus (Paradesmus) Beaumontii, Humbert & De Saussure, Verh. zool. bot. Ges.
Wien, vol. 19, p. 670.
1893 Orthomorpha beaumontii, Bollman, Bull. U.S. Nat. Mus., vol. 46, p. 196.
Probably misapplied:
1898 Prionopeltis Beaumontii, Attems, Denks. Ak. Wien, vol. 67, p. 357.
1937 Pratinus beaumontii, Attems, Tierreich, vol. 68, p. 122.
Material. — The label reads: “Polydesmus Beaumontii Gervais/M. Leguillou.
Bornéo.” (Mus. Paris), 1:9 (holotype).
Colour. — The colour has been described originally as being blackish brown,
270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
with the lateral keels paler. The specimens now has a dark chocolate brown colour,
the prosomites being a little paler. Lateral keels entirely pale dirty brown, tail also
pale. Ventral side pale brownish, sternites and legs dirty pale brownish.
Width. — 5.0 mm, the prosomites 3.6 mm.
Body segments. — Rather weakly constricted by a narrow waist, which is
distinctly longitudinally ribbed or “beaded” in the dorsal part down to the
level of the lateral keels, and smooth below that level. Prosomites somewhat dulled
by a fine cellular structure. Metatergites more shining, smooth or with some irre-
gular wrinkles in particular near the bases of the keels, hairless. Transverse furrow
well impressed, finely longitudinally striate, extending laterad to the bases of
the lateral keels. Transverse furrow present from the 5th segment to the 18th.
Sides up to the 4th segment rather densely but minutely granulate; from the 5th
Fig. 82—87. Orthomorpha beaumontii (Le Guillou) — 82: left side of the 2nd to the 5th
segments of the holotype ®, lateral view. 83: the same, dorsal view. 84: left side of the
10th and 11th segments of the holotype 2, dorsal view. 85: the same, lateral view. 86:
left side of the 16th to the 20th segments of the holotype 2, dorsal view. 87: anal segment
of the holotype 2, ventral view
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 271
segment onwards the granulation is weak to almost obsolete. Pleural keels well
developed, present from the 2nd to about the 15th segments. Up to the 4th
segment they are represented by a continuous ridge which is caudally produced
into a triangular lappet projecting behind the margin of the segment. From the
Sth segment onwards there is in each segment a rather weak swelling above the
anterior legs and a triangular lappet near the posterior border of the mecasomite.
Both, swellings and lappets are diminishing gradually in the subsequent segments
and disappear in about the 16th segment. The triangular lappets are projecting
very little behind the posterior margin of the segments up to about the middle of
the body.
Lateral keels. — (fig. 82—86). Strongly developed. Keels of 2nd segment
anteriorly rounded, somewhat shouldered at the base. Lateral border very weakly
rounded to almost straight. No tooth at the latero-anterior edge. Posterior edge
acutely angular and projecting rather strongly behind the margin of the segment.
Keels of 2nd segment scarcely below the level of those of the 3rd, turned slightly
ventrad. Marginal rim well developed, demarcated by a sharp premarginal furrow.
No ventral premarginal furrow. 3rd and 4th segments of subequal width,
somewhat wider than the 2nd. Keels of these segments very similar to those of the
2nd segment, but those of the 3rd somewhat less turned ventrad and those of
the 4th almost horizontal. 5th segment somewhat wider than the 4th. Keels from
the 5th segment onwards anteriorly distinctly shouldered at the base, anterior and
lateral borders widely rounded. Posterior edges acutely angular, projecting rather
strongly behind the posterior margin of the segments. Keels on a high level,
horizontal. Only in the 18th and 19th segments the keels are raised a little above
the horizontal level, but do not project above the mid-dorsal surface of the meta-
tergites. Poriferous keels slightly more prominent than the poreless. Marginal rim
rather thick dorso-ventrally, in poriferous keels scarcely thicker than in the
poreless, not abruptly widening near the pores. Dorsal premarginal furrow almost
reaching the waist. Posterior margin with a very fine rim. Lateral marginal rims
ventrally demarcated by a well developed furrow, present in poriferous and pore-
less keels. Pores lateral in a slight excavation of the rim, the excavation scarcely
visible from the dorsal side.
Sternite and legs. — Sternites in middle segments 11/,—11/, X as broad as
long, moderately densely set with rather short setae. Cross-impressions
moderately developed, the transverse furrow rather deeply and sharply impressed,
the longitudinal furrow less deep and rather wide. Legs of moderate length,
rather slender, moderately setiferous. Length of joints: 3>6>5>2>4>1. The
6th joint about two thirds of the length of the 3rd. The 5th joint about three
fifths of the 6th. Tarsal claw of moderate length.
Anal segment. — (fig. 87). Tail rather broad and thick at the base, of moderate
length. Sides moderately converging, scarcely concave. Basal lateral setiferous
tubercles moderately developed. Distal lateral tubercles and terminal tubercles al-
most obsolete. Dorsally near the base of the tail a shallow transverse depression,
ventral side of tail weakly concave. Valves with rather narrow rims, the setae on
weak tubercles. Scale triangular, the sides almost straight, the setiferous tubercles
well developed, projecting behind the posterior margin but not surpassing the
posterior edge of the scale.
De TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
Male. — Unknown.
Remarks. — That the type of O. beaumontii has turned out to be a female
specimen is certainly a rather unfortunate condition, since as a rule the specific
and generic identification of a Paradoxosomatid species is difficult if not impos-
sible with female characters.
Fortunately, however, O. beaumonti proves to belong to a group of species in
which the structure of the male gonopods is quite uniform and in which specific
characters lay mainly in the external structure.
Actual comparison of the type specimen with female co-types of O. weberi
(Poc.) has convinced me that weber? must be regarded as one of the closest
relatives of beaumontii. The entire morphology of the two species is quite similar,
in fact, they seem to differ only in the development of the lateral keels which in
weberi have the posterior edges a little less acute and distinctly less produced
caudad.
We may, therefore, safely assume that the gonopods of beaumontii ultimately
will prove to be of the type of those of weberi and related species like O. hydro-
biologica Att., O. francisca Att., O. karschii (Poc.), etc. (see group (I), p. 264).
Orthomorpha coarctata (Sauss.)
1860 Polydesmus (Paradesmus) coarctatus De Saussure, Mém. Soc. Genève, vol. 15, p. 297,
pl. 18, fig. 18.
1869 Polydesmus (Paradesmus) coarctatus, Humbert & De Saussure, Verh. zool.-bot. Ges.
Wien, vol. 19, p. 670 (1).
1937 Orthomorpha (Orthomorpha) coarctata, Attems, Tierreich, vol. 68, p. 62, fig. 75.
Previous records. — Borneo (1). Wide-spread in the tropical parts of all
regions.
Material studied. — East Borneo: Long Navang (Coll. Dr. E. MJÖBERG, Mus.
Stockholm), 15 &, 11 9. East Borneo: Songei Boh (Coll. Dr. E. MJÖBERG,
Mus. Stockholm), 5 ¢, 1.9. East Borneo: without nearer location (Coll. Dr. E.
MJÖBERG, Mus. Stockholm), 2 4. Maratua Id. off the East coast of Borneo
(coll. Dr. E. MJÖBERG, Mus. Stockholm), 5 4,3 9.
Remarks. — Not less than three generic names have been based on this species:
Asiomorpha Verh., 1939, Brasilogonopus Verh., 1944, and Orthomorphina Kraus,
1958, but at present I am not able to detect any reason for separating coarclala
either generically or subgenerically from O. beaumonti.
Oxidus Cook
1911 Oxidus Cook, Proc. U.S. Nat. Mus., vol. 40, p. 628.
1914 Orthomorpha (Kalorthomorpha) Attems, Arch. Naturg., vol. 80A, fasc. 4, p. 191.
1953 Kalorthomorpha, Attems, Mém. Mus. Nat. Hist. Nat. (n.s.), vol. 5, p. 163.
Type-species. — Fontaria gracilis C. Koch, by original designation. (Type-
species of Kalorthomorpha: Fontaria gracilis C. Koch, by original designation).
Range and number of species. — To be defined after a revision of the genus.
Remarks. — In its current concept, conforming with the subgenus Kalorthomor-
pha of ATTEMS in the Tierreich, this genus is at least as heterogeneous as the
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 273
previous one. The only species recorded from Borneo is certainly not congeneric
with Oxidus gracilis (Koch), but in the absence of material the definition of its
true taxonomic position is deferred to a later period.
“Oxidus” kuekenthali (Att.)
1897 Strongylosoma kiikenthali Attems, Abh. Senckenb. Ges., vol. 23, p. 484, pl. 21, fig. 9.
1937 Orthomorpha (Kalorthomorpha) kükenthali, Attems, Tierreich, vol. 68, p. 86, fig. 107.
Distribution. — Borneo. Celebes: Minahassa.
Remarks. — The occurrence of this species in Borneo as well as in Celebes
seems quite dubious. One of the records may be erroneous, and the species, there-
fore, may not pertain to the fauna of Borneo at all.
Euphyodesmus Att.
1931 Euphyodesmus Attems, Zoologica, Stuttg., vol. 30, fasc. 3/4, p. 125.
1937 Euphyodesmus (in part), Attems, Tierreich, vol. 68, pr 126:
Generic diagnosis. — 20 segments. Poreformula normal. Head without par-
ticulars; labrum tridentate. Antennae very long.
Segments rather strongly constricted. Prosomites with very fine cellular structure,
somewhat dull. Waist broad, distinctly and sometimes somewhat irregularly
longitudinally ribbed in the dorsal and lateral parts down to the level of the
stigmata. Metatergites shining, smooth or with some setiferous tubercles. Transverse
furrow well developed, rather deeply impressed, without longitudinal striation,
present from the collum onwards. Pleural keels present only in the 2nd, or in the
2nd and 3rd segments.
Lateral keels rather strongly developed, on all segments from the 2nd to the
19th provided with an acutely angular to subspiniform posterior edge surpassing
the posterior margin of the metasomites in most segments. Keels on a rather high
level, the posterior edges of those of the 5th and the 7th segments are turned
slightly upwards and project above the level of the metatergites. Keels of 2nd
segment only a little below the level of those of the 3rd segment. Marginal rim
of keels narrow, in poriferous segments widening only in the area of the pores.
Sternites much longer than wide, weakly setiferous, with well developed cross
impressions. Sternite of 5th segment of & with a strongly developed, distally
laminate process between the anterior legs. Sternites of the 6th, 7th and 8th
segments of the 4 without modifications. Legs long to very long. Those of the
first pair short and somewhat incrassate. Brushes of tibiae and tarsi present only in
a few anterior legs of the ¢, rapidly thinning out and disappearing.
Gonopod coxa of moderate length and rather thick, tapering a little towards
the distal end. Latero-distal surface with a setiferous area. Prefemur strongly
elongate, in one line with the femur, laterally and in part also medially well
demarcated from that joint, the demarcation almost in one line with the long-
itudinal axis of the femur. Femur rather weakly developed, straight, much shorter
than the coxa. Postfemur medially weakly to very sharply demarcated. Spermal
channel first running along the medial side of the femur, then curving towards
the anterior, lateral and finally posterior sides. Solenomerite of moderate length,
274 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
rather thick at the base, gradually tapering towards the distal end; the end either
simply acuminate or with an accessory lobe. Solenomerite arising from the
posterior side of the distal end of the postfemur; it is not sheathed by the tibio-
tarsus. Tibiotarsus arising from the anterior side of the distal end of the postfemur,
not distinctly demarcated from that joint, moderately developed or somewhat
reduced, consisting of a more or less compound or of a simple lamella. Lamina
medialis or lamina lateralis both not developed.
Type-species. — Euphyodesmus gracilis Att., by original designation.
Range. — Borneo.
Number of species. — Three.
Remarks. — After the reallocation in the genus Pratinus Att. of Euphyodesmus
greeni Att. and E. vector Chamb. in the preceding pages of this paper, a single
species was left in Ewphyodesmus. From the present collection two new species
are added.
The anomalous structure of the gonopods in Exphyodesmus makes a homolo-
gization of the distal parts of the acropodite rather difficult. Doubtless, the
aberrant course of the spermal channel in the femoral division has been caused by
a torsion of about 180° of the distal part of the acropodite as against its basal
part. The true nature of the parts distad of the femur, however, is less easily
determined and, in fact, could be ascertained only after the examination of the
gonopods of the newly described E. postfemoralis.
As will be seen in the gonopod drawings for E. gracilis (fig. 94—95), the
acropodite has a distinct constriction somewhat proximad of its branching into
solenomerite and tibiotarsus, a constriction which laterally is accompanied by a very
thin laminate transverse crest and which is the only indication of an articulation in
the acropodite.
Comparison of this with the usual type of gonopods in the Oriental Para-
doxosomatidae might lead to the conjecture that the constriction represents the
boundaries beween the tibiotarsus on the one hand and the femur and postfemur
combined on the other. However, this would imply that the solenomerite in
Euphyodesmus arises from the base of the tibiotarsus, a quite unusual condition
since it is generally admitted that the solenomerite in the Paradoxosomatidae is a
process arising from the postfemur, or, if this joint is not demarcated, from the
distal end of the femur.
Indeed, a comparison of the gonopod drawings of E. postfemoralis with those
of gracilis shows that the constriction in the acropodite of the latter species marks
the end of the femur proper. The undivided part of the acropodite distad of the
constriction therefore must be regarded as a postfemur. In postfemoralis there is a
sharp demarcation between femur and postfemur on the medial side, homologous
to a similar demarcation visible on the lateral side of the acropodite of the gono-
pods of, e.g., Arthrogonopus and Gigantomorpha. In postfemoralis the postfemur
is furthermore also more clearly demarcated from the tibiotarsus, at least laterally,
and, medially from the solenomerite.
Thus, it is interesting to note that in Euphyodesmus, where the demarcation
between postfemur and tibiotarsus tends to disappear, the demarcation between
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 275
femur and postfemur, which otherwise has disappeared in so many of the Para-
doxosomatidae, remains distinct.
It is necessary to emphasize here, that I have considered the possibility that what
is regarded here as a solenomerite could have been a solenomerite concealed in a
closely applied solenophorous part of the tibiotarsus. Such a condition, of course,
would involve a quite different homologization of the distal part of the acropodite.
However, by studying a preparation of a gonopod crushed somewhat by the cover
glass I could ascertain that the solenomerite is a simple, undivided process.
The fact that both ATTEMs and CHAMBERLIN have referred species of the
genus Pratinus to Euphyodesmus might suggest a close relationship between the
two genera. In reality, however, there exists only a superficial resemblance. In
particular the gonopods of Pratinus are fundamentally different from those of
Euphyodesmus. In Pratinus the spermal channel follows a straight course along the
medio-anterior side of the femur, the solenomerite arising, consequently, from
the anterior side of the distal end of the femur. Moreover, the tibiotarsus in
Pratinus seems to be distinctly demarcated from the femur, and a postfemur is
apparently not marked.
In the morphology of the body segments, in particular in the shape of the
lateral keels Euphyodesmus is also quite distinct from Pratinus. The subdorsal
location of the pores and the more or less rhomboid dilatation of the marginal
rim of the poriferous keels are strongly reminiscent of similar structures found in
some species of Tectoporus, e.g., T. hispidus Jeek., 1951.
Key to the species. — The species of Euphyodesmus are very similar in their
external morphology. Important specific differences are to be found mainly in
the male gonopods.
1. Metatergites with two transverse rows of four granules each; in the anterior
rows the granules are setiferous. Tibiotarsus of the gonopods simply laminate.
Ends ongsolcnomenitegbifurea te Serra E. granulifer spec. nov.
— Metatergites without granules, hairless. Tibiotarsus of the gonopods more
complieated. Endrotssolenomerite notibitureate nn 2
2. Postfemur of gonopods medially sharply demarcated from the femur by a
furrow. Solenomerite distinctly longer than tibiotarsus, the end with a triangular
lamellar lobe. Head much wider than collum ...... E. postfemoralis spec. nov.
— Postfemur of gonopods medially not sharply demarcated from the femur.
Solenomerite about as long as tibiotarsus, simply acuminate at the end. Head
pogmuch dentino UM E. gracilis Att.
Euphyodesmus gracilis Att.
1931 Euphyodesmu: gracilis Attems, Zoologica, Stuttg., vol. 30, fasc. 3/4, p. 126, fig.
195—199.
1937 Euphyodesmus gracilis, Attems, Tierreich, vol. 68, p. 127, fig. 163—164.
Previous record. — West Borneo: Sungei Malang.
Material studied. — West Borneo: Bungan River, July, 1894 (Borneo Exped.,
Coll. Dr. A. W. NIEUWENHUIS, Mus. Leiden), 1 4, 1 9.
Colour. — Head brownish yellow to yellowish brown. Antennae pale brown,
276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
C. A. W. JEEKEL : Paradoxosomatidae from Borneo Di
darker in the distal half of the 6th joint and in the 7th joint. Distal end of the
7th and the 8th joint whitish. Collum and subsequent segments castaneous, the
latero-posterior edges of the collum and of the lateral keels of the body segments
paler, yellowish. Venter pale brownish, the sternites yellowish. Legs proximally
and distally yellowish, the middle part from the distal end of the femur to
about the middle of the tarsi dark castaneous. In the two last pairs the dark
colour extends to the base of the femur. Anal segment castaneous, its ventral side
paler.
Width. — & 1.6 mm;:9 1.9 mm.
Head and antennae. — Labrum rather widely but weakly emarginate. Clypeus
weakly convex, weakly impressed towards the labrum; the lateral margins widely
and rather weakly emarginate. Surface of head smooth and shining, densely to
rather densely setiferous in the clypeal and frontal parts. Vertex with four setae.
Antennal sockets separated by little more than the diameter of a socket or by
about one third of the length of the 2nd antennal joint. Postantennal groove
well developed, the wall in front of it rather prominent. Vertex strongly convex,
not demarcated from the frontal area. Sulcus weakly impressed, not reaching the
upper level of the antennal sockets. Antennae (fig. 88) slender. Length of joints:
3>4>5 = 2>6. The 6th joint about half as long as the 3rd. Joints of subequal
width, only the 6th somewhat thicker than the others. Pubescence of antennae
moderate to, distally, rather dense.
Collum. — (fig. 89—90). Distinctly narrower than the head, subsemicircular
in dorsal outline. Anterior border evenly rounded. Posterior border widely and
weakly emarginate in the middle, almost straight laterally. Latero-posterior edge
narrowly rounded, obtusely angular. Surface rather shining, somewhat irregularly
uneven, apparently hairless. At about one third from the posterior margin a well
developed transverse furrow. Surface of collum transversely almost flat in the
middle, much more convex towards the lateral sides; the lateral sides slightly
raised but not attaining a horizontal level. Marginal rim narrow and rather weakly
demarcated, present only along the latero-anterior border.
Body segments. — Metatergites somewhat uneven in a few anterior segments,
but in most segments smooth with a few weak wrinkles. Transverse furrow
present up to the 18th segment, from the 5th to the 17th segments running laterad
to the base of the keels and curving caudad there to end at the posterior margin of
the keels. Metatergites hairless, only the 19th segment with a transverse row of
four hairs. Sides up to the 4th segment rather densely granulate, smooth or weakly
granulose from the 5th segment onwards. Pleural keels very weakly present in
the 2nd segment only.
Lateral keels. — (fig. 8992). 2nd segment a little wider than the collum.
The border of the keels anteriorly rounded, laterally straight with two very weak
Fig. 88—95. Euphyodesmus gracilis Att. — 88: antenna, without the basal joint, of the 4.
89: left side of the head and the three anterior segments of the 4, lateral view. 90: the
same, dorsal view. 91: left side of the 10th and 11th segments of the @, lateral view.
92: the same, dorsal view. 93: leg of the 7th segment of the &. 94: right gonopod of
the 4, medial view. 95: distal portion of the telopodite of the left gonopod of the 4,
lateral view
278 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
notches. Posterior edge acutely angular, but not sharply pointed; not projecting
behind the margin of the segment. Posterior border emarginate. Marginal rim
weakly developed, obsolete in the posterior half of the keels. 3rd and 4th segments
of subequal width, scarcely narrower than the 2nd. Keels subsimilar to those of
the 2nd segment; the notches of the lateral border vestigial or absent. The
posterior edges are more acute, and project a little behind the margin of the
metasomite in the 4th segment. Up to the 4th segment the lateral keels are
horizontal. 5th segment a little wider than the 4th. Lateral border of the keels
from the 5th segment onwards with widely to very widely rounded latero-anterior
borders which laterally are straight or even a little concave in poriferous segments.
Latero-posterior edges very acute and sharply pointed, in all segments projecting
behind the posterior margin of the metasomites. Posterior borders of keels
emarginate. Lateral margins often with a weak notch. Keels raised a little above
the horizontal level. The posterior edges of those of the 5th and 7th segments bent
upwards and projecting dorsad of the level of the metatergites. Posterior edges of
the keels of the 13th, 15th, and 17th segments curving a little ventrad. Poriferous
keels extending slightly more laterad than the poreless. Marginal rim of poreless
keels narrow but distinct. The premarginal furrow running from near the waist
to near the posterior margin. In poriferous keels the premarginal furrow reaches
the posterior border, and the rim widens considerably in the area of the pores.
Pores latero-dorsad, situated in a wide and shallow excavation of the rim.
Posterior margin of all keels with a fine marginal rim. On the ventral side of the
keels from the 5th segment onwards the marginal rim is demarcated by a pre-
marginal furrow in the middle part of each keel.
Sternites and legs. — Sternites of middle segments somewhat more than two
times longer than broad, sparsely setiferous. Cross impressions represented by a
deep transverse depression and a moderately deep longitudinal impression. Sternal
process of the 5th segment narrow, subrectangular, about two times longer than
broad; the distal end widely and weakly rounded. The process is directed ventrad
and does not project before the anterior margin of the sternite. Anterior side
distally with a small brush of moderately short setae, normally setiferous.
Posterior portion of the sternite of the 5th segment normal. Legs (fig. 93) long;
the last two pairs distinctly longer than the preceding pairs. First pair of legs
rather short, distinctly incrassate; the joints of subequal length, but the tarsus
much longer. The first pair has well developed tibial and tarsal brushes, which
in a few of the following pairs are rapidly thinning out and disappear. Legs of
middle segments rather weakly to, distally, moderately setiferous, especially on
the ventral side of the joints. Length of joints: 3 = 6>5>4>2>1. The 5th
joint about two thirds of the 6th.
Anal segment. — Tail of moderate length. Sides rather strongly converging,
straight. The end narrowly truncate, rounded, with weak lateral and terminal
tubercles. Valves with rather narrow and rather low marginal rims. Scale
triangular; the sides rounded. Setiferous tubercles small, not projecting.
Gonopods. — (fig. 94—95). Postfemur indicated by a constriction and by a
thin transverse lateral crest. Solenomerite simply acuminate, without secondary
processes. Tibiotarsus moderately developed, rather complicated by several lobes,
apparently supporting though not actually sheathing the solenomerite.
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 279
Fig. 96—98. Euphyodesmus postfemoralis spec. nov. — 96: left side of the head and the
collum of the holotype ¢, dorsal view. 97: right gonopod of the holotype &, medial view.
98: distal end of the left gonopod of the holotype &, lateral view
Female. — Distinctly more robust than the male. Aside from the usual secondary
sexual characters also differing in that the somites are somewhat less strongly con-
stricted in the waist area. The waist a little narrower. Sternites of middle segments
about 13/, longer than broad. First pair of legs incrassate as in the male,
but without dense brushes. The other legs comparatively a little shorter than in
the male.
Remarks. — Although there can be little doubt as to the correctness of the
identification of the present material with graczlis Att., there are some minor dif-
ferences between the gonopods of the male at hand and the type specimen described
by ATTEMS. The various lamellae of the tibiotarsus differ slightly in their outline,
whereas the solenomerite is curved a little in the end instead of being almost
straight. No doubt these variations are at most of a subspecific nature, but of not
sufficient significance to justify a denomination.
Euphyodesmus postfemoralis spec. nov.
Material. — East Borneo: Mt. Tibang, 1700 m (Coll. Dr. E. MJOBERG, Mus.
Stockholm), 5 4 (one of which the holotype), 4 @, 2 juv. 9. East Borneo: Mt.
Tibang, 1400 m (Coll. Dr. E. MJÔBERG, Mus. Stockholm), 6 &, 3:9, 2 juv. 4,
6 juv. @. East Borneo: Mt. Tibang, 1300 m (Coll. E. MJÖBERG, Mus. Stock-
280 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
holm), 2 9, 3 juv. &, 2 juv. 2. East Borneo: without nearer location (Coll. Dr.
E. MjÖBERG, Mus. Stockholm), 2 4,1 2.
Differing from gracilis in the following characters.
Colour. — Most of the material of this species apparently was not in full
colour when it was collected. The colour of the darkest specimens, however, agrees
with that of gracilis, but the legs almost wholly lack the dark colouring.
Width. — Holotype & 1.6 mm. The other & in the order of the above
enumeration; 1.6 mm, 1.5 mm, 1.5 mm, 1.4 mm; 1.5 mm, 1.5 mm, 1.5 mm,
1.4. mm, ls saran Ib) on Ice, Joby sag Oe 2,2 ANW 22 foon, Zlk ren,
2.0 mm; 2.0 mm, 2.0 mm, 1.8 mm; 2.0 mm, 2.0 mm; 2.1 mm. Juvenile 4
with 19 segments: 1.0 to 1.1 mm. Juvenile © with 19 segments: 1.0 tot 1.4 mm.
Collum. — Much narrower than the head, as a consequence of the strongly
inflated lateral sides of the latter (fig. 96).
Gonopods. — (fig. 97—98). Femur sharply demarcated from the postfemur
by a furrow on the median side. Solenomerite more strongly developed, the end
with a triangular lamellate lappet. Tibiotarsus shorter than the solenomerite,
apparently free from the solenomerite.
Female. — Differing from the 4 in the same way as in gracilis.
Juvenile. — Although in general features agreeing with the adult 9 specimens,
the juvenile specimens have much more “normal” aspect because of the weaker
constriction of the waist area of the body segments. The antennae and legs are
comparatively much shorter than in the adults. The posterior edges of the lateral
keels are less produced caudad.
Euphyodesmus granulifer spec. nov.
Material. — East Borneo: Pajan River (Coll. Dr. E. MJOBERG, Mus. Stock-
holm), 4 4 (one of which the holotype), 2:2, 3 juv. &, 1 juv. 9. East Borneo:
Mt. Tibang, 1400 m (Coll. Dr. E. MJÔBERG, Mus. Stockholm), 1 9. East
Borneo: without nearer location (Coll. Dr. E. MJOBERG, Mus. Stockholm), 1 ¢.
Differing from gracilis in the following characters.
Colour. — The best preserved specimens of this species are darker than
gracilis. Head dark castaneous, the clypeal area lighter, yellowish towards the
labrum. Antennae also dark castaneous, only the tip of the 7th and the 8th joints
whitish. Collum and subsequent body segments dark castaneous, but the collum
with a dirty whitish zone along the anterior margin and similarly coloured lateral
edges. Lateral keels of the 2nd to the 4th segments entirely, of subsequent seg-
ments only in the latero-posterior portion pale yellowish. Dark portion of the legs
also deeper brown than in gracilis, the brown colour extending to the end of the
tarsi.
Width. — Holotype ¢ 1.6 mm. The other & in the order of the above
enumeration: 1.6 mm, 1.5 mm, 1.5 mm; 1.4 mm. ‘9 : 2.0 mm, 1.9 mm; 1.9 mm.
Juvenile 4 with 19 segments: 1.4 mm, 1.3 mm, 1.2 mm. Juvenile 9 with 19
segments: 1.3 mm.
Head and antennae. — Antennal sockets separated by about 4/5 of the
diameter of a socket, or by a quarter of the length of the 2nd antennal joint.
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 281
Fig. 99— 100. Euphyodesmus granulifer spec. nov. — 99: right gonopod of the holotype 4,
medial view. 100: distal end of the left gonopod of the holotype &, lateral view
Vertigial sulcus running downward to just below the upper level of the antennal
sockets.
Collum. — 4 + 4 minute but distinct setiferous granules along the anterior
margin. The hairs of moderate length, rather stiff.
Body segments. — Metatergites with a row of 4 setiferous granules behind the
waist, and 4 somewhat less distinct hairless granules behind the transverse sulcus.
Waist with weakly developed, somewhat irregular ridges. Pleural keels on the
2nd segment represented by a triangular lappet projecting downward. In the 3rd
segment weakly developed, in subsequent segments absent.
Lateral keels. — 2nd segment slightly narrower than the collum, the keels
pointed. In general the lateral keels are slightly more strongly developed than in
the two preceding species, the posterior points being a little longer.
Gonopods. — (fig. 99—100). Postfemur demarcated from the femur by a thin
transverse crest on the medial side. Solenomerite bifurcate in the end. Tibiotarsus
a simple subrectangular lamina, entirely free from the solenomerite.
Female. — Differing from the ¢ in the same way as in gracilis.
Juvenile. — See the remarks made for the juvenile of postfemoralis.
“Strongylosoma” nodulosum Att.
1897 Strongylosoma nodulosum Attems, Abh. Senckenb. Ges., vol. 23, p. 486.
Distribution. — Borneo.
Remarks. — The male characters of this species are unknown, reason why its
282 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 6, 1963
generic position has remained a mystery. Even in the light of the present additions
to the Borneo Paradoxosomatid fauna little can be said of its relationship. The
weak development of the lateral keels at least seems to exclude the possibility
that it might belong to Gigantomorpha, Orthomorpha or Euphyodesmus. The
colour pattern as described by ATTEMS suggests a similarity with the colour of
Borneonina retrorsa. To say more would be mere speculation, so nodulosum is
condemned to remain an enigma probably for a long time to come.
REFERENCES
ATTEMS, C., 1897. Myriopoden. Abh. Senck. Naturf. Ges., vol. 23, p. 473—536, pl. 21—24.
1898. System der Polydesmiden. I. Theil. Denkschr. Ak. Wiss. Wien, vol. 67,
p. 221—482, pl. 1—11.
—, 1914. Die indo-australischen Myriopoden. Arch. Naturg., vol. 80A, fasc. 4, p.
1398 jelly) =).
—, 1931. Die Familie Leptodesmidae und andere Polydesmiden. Zoologica (Stuttg.),
vol. 30, fasc. 3/4, p. 1—150.
——, 1936. Diplopoda of India. Mem. Ind. Mus., vol. 11, p. 133—323.
——, 1937. Polydesmoidea I. Fam. Strongylosomidae. Tierreich, vol. 68, p. I—XXII,
I—300.
—, 1953. Myriopoden von Indochina. Expedition von Dr. C. Dawydoff (1938—
1939). Mém. Mus. Nat. Hist. Nat., new ser., vol. 5, p. 133—230.
BOLLMAN, C. H., 1893. The Myriopoda of North America. Bull. U.S. Nat. Mus., vol. 46, p.
1—210.
CARL, J., 1912. Die Diplopoden-Fauna von Celebes. Rev. Suisse Zool., vol. 20, p. 73—206,
pl. 5—6.
, 1941. Orientalische Polydesmoiden. Rev. Suisse Zool., vol. 48, p. 359—376.
CHAMBERLIN, R. V., 1941. New Polydesmoid Diplopods intercepted at Quarantine. Proc.
Ent. Soc. Wash., vol. 43, p. 32—35.
1945. On some Diplopods from the Indo-Australian Archipelago. Amer. Mus. Nov.,
Nr. 1282, p. 1—43.
Cook, O. F., 1895. East African Diplopoda of the Suborder Polydesmoidea, collected by
Mr. William Astor Chanler. Proc. U.S. Nat. Mus., vol. 18, p. 81—111, pl. 2—6.
——, 1911. The Hothouse Milliped as a new Genus. Proc. U.S. Nat. Mus., vol. 40, p.
625—631.
GERVAIS, P., 1847. Myriapodes, in: WALCKENAER & GERVAIS, Histoire Naturelle des Insectes
Aptères, vol. 4, p. 1—133.
HUMBERT, A., & DE SAUSSURE, H., 1869. Description de divers Myriapodes du Musée de
Vienne. Verh. zool.-bot. Ges. Wien, vol. 19, p. 669—692.
JEEKEL, C. A. W., 1951. The genus Tectoporus Carl (Diplopoda, Polydesmoidea, Strongylo-
somidae). Ent. Ber., vol. 13, p. 266—271, 281—285.
1953. Two new Strongylosomidae from Indochina (Diplopoda, Polydesmida).
Beaufortia, vol. 2, fasc. 29, p. 1—8.
Kraus, O., 1958. Myriopoden von den Galapagos-Inseln. Senck. biol., vol. 39, p. 97—102.
Pocock, R. I., 1895. The Myriopoda of Burma, Pt. IV. Ann. Mus. Genova, vol. 34, p.
787—834.
, 1895—1910. Chilopoda and Diplopoda, in: Biologia Centrali-Americana, p. 1—
Zap
DE SAUSSURE, H., 1859. Note sur la famille des Polydesmides, principalement au point de
vue des espèces américaines. Linn. Ent., vol. 13, p. 318—327.
—, 1860. Essai d'une faune de Myriapodes du Mexique, avec la description de quelques
espèces des autres parties de l’Amerique. Mém. Soc. Phys. Genève, vol. 15, p.
1—133, pl. 1—7.
SILVESTRI, F., 1896. I Diplopodi. Parte I. — Sistematica. Ann. Mus. Genova, vol. 36, p.
121—254.
>
2
>
C. A. W. JEEKEL : Paradoxosomatidae from Borneo 283
VERHOEFF, K. W., 1939. Zur Kenntnis ostasiatischer Diplopoden III. Zool. Anz., vol. 127,
p. 113—125.
——., 1941. Studien an äthiopischen Diplopoden. Jena. Z. Naturw., vol. 73, p. 231—274,
pl. 10—13.
——, 1944. Ueber einige Diplopoden aus Minas Geraes (Brasilien). 146. Diplopoden-
Aufsatz. Arq. Mus. Nac. Rio de Janeiro, vol. 37, p. 249—288, 8 pls.
AT
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TIJDSCHRIFT
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AFRICAN SPECIES OF THE GENUS BACTRA STEPHENS
(LEPIDOPTERA, TORTRICIDAE)
BY
A. DIAKONOFF
Rijksmuseum van Natuurlijke Historie, Leiden
After having published surveys of the species of the genus Bactra Stephens from
tropical Asia (1950, 1956), and from the Palaearctic and Mediterranean Regions
(1956, 1959 and 1962), I now present a preliminary survey of the species from
the Aethiopian Region, chiefly South Africa.
Does the genus show a rich speciation in tropical Asia, in South Africa this
abundance is almost exceeded. At first I had the impression that India might
represent the country of origin, a zoocentre, of Bactra species. The present results,
however, suggest that we may be dealing with a very old group which may have
originated in some ancient central region of the Old World Tropics, somewhere
between the Asiatic and African continents, to spread east and west. This might
explain the partly very puzzling distribution of certain species groups, e.g., the
boschmai group: in New Guinea, Africa and the West Indies (!), the coronata
group: in Australia, South Asia, and South Africa, etc.
The species of Bactra can easily be grouped in five natural subgenera, four of
which have been defined by me previously (1956), while the fifth is described
below. The general character of these subgenera, their taxonomic ‘behaviour’, is
not at all similar. So the species of the most extensive subgenus, Chz/oides Butler,
possess strongly specialized genitalia in the two sexes and show marked differences
from one another. The group offers the greatest diversity of genital structure which
makes its study very fascinating. Chiloides contains many species occurring
throughout the tropical and subtropical regions of the Old and the New World.
These features suggest a great antiquity of the group.
On the other hand, the two following subgenera, Bactra and Nannobactra, are
different from Chiloides, but are rather similar to each other with regard to this
taxonomic behaviour, viz., within each subgenus the species show but minor dif-
ferences of genitalia; especially the females are rather uniform in each of these
two subgenera. While those of Bactra Stephens may mostly be discriminated
without difficulty, the females of Nannobactra Diakonoff often are so similar
that their separation becomes a hazardous or even an impossible task. These two
subgenera must be much younger than Chiloides, although of unequal origin:
while Bactra must have developed from some simplified Chiloides-like ancestors
through the loss of the valvula, Nannobactra may be regarded as a quite different
off-shoot of the Chzloides stock, having originated through the speciation of the
cucullus portion of the valva and of its armatures, but with the retention of the
valvula.
285
286 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
Finally Spinobactra subgen. nov. and Noterazla (New Zealand) are endemic
developments of Bactra-like ancestors.
The rather numerous species are very similar in appearance and facies. Most
are subject to considerable variation along parallel lines, so giving rise to a series
of infraspecific forms which in diverse species look very much alike. Therefore,
the student is almost entirely thrown on the study of the genital characters —
and there he finds a remarkable richness of structure, in males as well as in females.
These structures are of a great specific rigidity and have a pronounced specific,
as well as subgeneric, taxonomic value.
EDWARD MEYRICK, who described most African Bactra species (1909, 1911),
and a few other Lepidopterists of his time (ZELLER, 1852, WALKER, 1866, WAL-
SINGHAM, 1881, MABILLE, 1900), were not aware of this internal diversity and
used exclusively external characters. The result was rather chaotic.
So the extensive collection of Bactra species in the Transvaal Museum, forming
the backbone of this survey, has been named by MEYRICK for more than three
quarters. The results bear evidence of the impossibility to discriminate these
species without dissecting and mounting the genitalia of every available specimen.
Therefore, the older identifications be better entirely disregarded, except, of course,
the types. That is the reason why I am not citing any of MEYRICK's determination
labels in most cases.
In order to facilitate the use of some terms in the male genitalia in the subgenus
Bactra, the reader is referred to fig. 1. The new concept, defined below and
depicted in that figure is the ‘’antebasal hairs’. For explanations of a few more
new terms I may refer to the descriptions of the subgenera.
The present survey is preliminary, for I expect to find more interesting material
of African Bactra in other museums on that continent and elsewhere, for which I
did not apply as yet. Not to delay the present results I am publishing them now,
although several problems had to remain unsolved, chiefly by lack of sufficient
material.
Two subgenera and 17 species are described as new and 9 earlier described
species are recorded.
For material and information I am greatly indebted to the following persons
and institutions. In the first place, to Dr. L. Vari, Transvaal Museum, Pretoria,
South Africa; furthermore to Mr. J. D. BRADLEY, British Museum (Natural
History), London, England; Dr. F. Kasy, Naturhistorisches Museum, Vienna,
Austria; Dr. J. F. GATES CLARKE, U.S. National Museum, Washington, D.C.,
U.S.A.; Dr. P. E. L. VIETTE, Muséum National d'Histoire Naturelle, Paris, France;
Mr. E. TAyLoR, Hope Department, University Museum, Oxford, England, and
finally, to the Information Attaché, South African Embassy, The Hague, Nether-
lands, for information on localities.
Figures 7 and 8 were made by W. BERGMANS, 38 and 73 by Miss R. VAN
CREVEL, 64 and 72 by A. VAN Dijk, the rest by myself.
KEY TO THE AFRICAN SPECIES OF Bactra BASED ON GENITAL CHARACTERS
Males
I A,'separate valvulawabsenta.., i e 2 Ae RO A CRN
10.
A. DIAKONOFF : African species of Bactra 287
A well-developed, clavate and coronate valvula present between cucullus
andısacaulust m nu e AIN MES GAI O)
. Entire cucullus beset na numerous rigid pce sil à not sous, with
an edge of very dense spines, punctulate area extending over the rest of
sacaulus rn un: 5 . +. + . Spinobactra spinosa
Cucullus bristles and Spies demen only dons its ventral portion; sacculus
bulbous, without a dense fringe of spines, with Spc, only. Subgenus Bactra 3
. Bases of socii with a as sclerotized nice making the socius to stand
UDHODE EE oe a lanceolana
Bases of socii never Sint a cad re socius oh Bending or recumbent 4
. Punctulate area of sacculus extended, anne its distal half with numerous
spinesse zur. 5 6 Wot sd ee JACEE
Punctulate area ttc most tieel as Ess HUMETOUS KIN MERENS
SEMISWSELIESMLOEMEHOLNStLOD EPS PINES iy) AAN ON Nico pa
Ms series formed of small bristles or hairs. . . DER SEN. 6
. Uncus long and slender, pointed and curved, thus corona of spines long.
Sacculus small, rigid, triangular or quadrate in profile, Spc, dense graminivora
Uncus of normal length. Sacculus pani spherical, not rigid, nor sclerotized;
SPC mMoLmal sisi a. Man 7
. Terminal patch of cucullus ride: not developed chee PAIE do not extend
basad byond the edge of sacculus . . . . MC LL
Terminal patch of cucullus bristles well- developed extending basad beyond
the edge ofgisacculust o, (IE LN aS
. There is a well-defined, Li Jas wart ay on edge of
sacculus and punctulate area, beset with spines and hairs. . . scrupulosa
No such isolated, elevated wart on sacculus . . . . ran beg
. Punctulate area semioval, well-defined, longer; Spc, does Senden straight
and longer; basal hairs normal, not extended; terminal patch extended basad;
its anterior edge forming a well-defined elevated ridge, beset with spines.
Species with broader, less pointed wings and a straight termen . . confusa
Punctulate area shorter, narrowed against top, at base not limited but extended
transversely and merging into patch of basal hairs; Spc, spines less numerous
(3—4), shorter, flattened and curved; terminal patch less extended basad,
its anterior edge not elevated. Species with dilated, pointed fore wings and
Agsinuate termenw rar a ot Mia
Cucullus diversely shaped, ngi) donde not bee are with nor-
mal spines and bristles. Subgenus Chzloides . . . . CA Bike TT
Cucullus short and broad, rounded and sclerotized, Sio one to three series of
very large spines, increasing in size downward. Subgenus Nannobactra . 23
. Vinculum triangularly thickened in middle, dentate. . . . . venosana
Vinculum smooth, never dentate . . . Hove ae 12
. Base of sacculus outwardly and inwardly ch Beels dense long spines
SUT a ARA ENT A PEREN o DI TECA kc PI NO SI punctistrigana
If sacculus spined then not only at base and not so densely. . . . . 13
. Sacculus with a marginal corona of long spines of equal length. . . 14
Sacculus without a marginal corona of equal spines (seldom a few spines of
288
26.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
diverse length at the bottom ally, erispinesishort) 2. Re nn LG
. Top of sacculus strongly extended laterad, so as to form a naarden process
erowneds with longsspinesg (ed tornastı ap) nn DE
Sacculus broadly rounded, circular or semicircular in circumference . . 15
. Juxta very long, almost twice as long as sacculus is broad . . . sinassula
Juxta of normal length, not longer than sacculus is broad. . . . fasciata
. Top of very concave sacculus with an inside-turned process. . . 017
Top of sacculus sometimes with a low and obtuse ridge but Ao € a process
. Apical process of sacculus long, conspicuous, Ance TEE or
Apical process of sacculus aculety-triangular, more or less concealed . aletha
. Aedeagus short, straight, tubular, not exceeding width of juxta. Sacculus
with not numerous stout or thin spines. . . o 19
Aedeagus long, mostly curved, by far exceeding ill af hea if moder
then flattened laterally Ro 22 2 me 2 IKC NP er 20
MSacculus with 1l0S touts Pine EN MEN NC AEEA
Sacculus with a few (4—5) thin spines. . . nilo ie o ane
. Aedeagus moderate, strongly flattened tica arenden very broad, with
denticulations before middle of upper edge. . . . . . . stagnicolana
Aedeacusmlong;mnevermlattened laterally en Sr re
SySacculuspmotispined RE EA 01474212
Sacculus distinctly spined . . . . . Ut Rte eee
. Sacculus spines long and slender, of ess ont, ne in patches
trimera
Spass pires shar numerous 6 alone lover half sé outer alles, and few on
{0 PRONES ACCUSE u. toa . . » = thabdonoma
. Valvula extremely large, corona rie a Tony & sinuate band . . sardonia
Valvula of normal size . . . Wte Suter ER
. Tegumen with rounded- rer shoulder lobes Les of gnathos)
distinctly aciculate . . . B à CAMO
Tegumen either with entirely ora ti: leken, or these lobes obliterate
25
Shape en zen a coi its upper al Gone by a small and
globular punctulate area, lower half appearing du spines of punctulate
area numerous, long. . . no ee LORE
Sacculus not thus vertically stonde punctulate area En its larger part,
its) lower |portion notithus extended:#spiness hote NE 26
Spines of punctulate area numerous, dark, moderate, pointed; corona spines
inequal: ultimate spine spindle-shaped, thick, point slightly bent. . minima
Sacculus spines few, short, less pointed, not conspicuously dark; corona spines
subequal: two or three ultimate spines approximately similar in size and
shape) an. mod nuit HOLMAN ae lie Swear er Ware Guild
Females
. No or very faint sclerotizations of the seventh segment; eighth + ninth seg-
ment not sclerotized; if posterior edge of tergite and bases of anapophyses
A. DIAKONOFF : African species of Bactra 289
slightly sclerotized, and there are small angulate sclerites on both sides of the
ostium, then signum is absent. Subgenus Bactra (part.) . . . . . . 2
Eighth + ninth segment more or less distinctly sclerotized, sometimes strongly
sclerotized and modified. Signum usually larger, basket-shaped, always present
Ostium moderate, pear-shaped, lamella postvaginalis vertical, marrow and
wrinkled, posterior edges of genital sternite also finely wrinkled. Ostium
flanked by two weak oval plates, their median edges thickened, so as to form
a small refracting crescentic structure (figs. 6—7). Colliculum tubular, short.
Signum absent. Subgenus Bactra (part.) . . . Spy onia
Eighth + ninth segment diversely sclerotized, usually only along its edge, or
only sterigma moderately sclerotized, or there is a pair of rounded sclerites
at the sides of sterigma; the latter always weakly modified, ostium usually
being surrounded by aciculate or minutely haired simple folds or rounded
prominences; genitalia rather uniform , species difficult to separate. Subgenus
Nannobactra re zi
. Lamella postvaginalis tn a transverse Toron band at Ane ioni
or with a series of horizontal fine folds, or almost simple, never cap-shaped
Lamella postvaginalis with a more or less developed cap-shaped structure,
concave below. (Ostium su on top of a prominence, visible only
inepro file) Mn U 4
. Lamella postvaginalis el a series of ie horizontal mer straight folds,
not concave below, sometimes hardly traceable. Ostium, shallow, of a crescentic
aspect, its outer edge concave. Lobus analis Be with a gradu-
ally rounded top, outer edge convex. . . . . graminivora
Lamella postvaginalis with a well-defined transverse Gand of fine punctul-
ations, edged above and beneath by a smooth fold, dilated and truncate at
extremities. Lobus analis not distinctly semioval, its outer edge not so convex,
top subobtuse but not gradually rounded. . . . . . . . lanceolana
. Lamella postvaginalis with a simple cap, being a slender curved fold over
ostium; rarely minute omg present above this. a always present,
althouch@ smaller rn . . furfurana
Cap of lamella le more dedalo or are are boni folds at
the sides of the sterigma, or some sclerotizations, or there is more punctulation
above the cap . . 5
. Apophyses very biz Fico DE of nose para rei langs dan lobus
analis. Lamella postvaginalis with punctulations and also several bristles on
basal warts. Signum absent . . . ne alte CU
Apophyses normal, free part of pese Abone as slong as lobus analis. No
bristles on lamella postvaginalis . . . . hentia 6
. Cap of lamella postvaginalis broad, formed of one or two ) Loro median
folds and a small additional cap on each side. Ostium wider. A more or less
sclerotized curved plate on each side TRE ostium. Signum absent .
confusa
Ce af cele posse: ‘ex proach hau, ritor lateral caps;
15:
14.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
punctulation above this cap distinct, forming a well-defined elongate field;
no plates at the sides of sterigma, instead a pair of darkly coloured small rods.
Rostral margin of genital tergite often sclerotized and appearing as a semi-
annular structure joining the bases of anapophyses. Signum absent . tradens
Cap of lamella postvaginalis small, very narrow in middle, with slight lateral
caps, the lateral edges traceable downwards and forming some additional folds
on each side of sterigma; there is Voi some punctulation above cap. Signum
ABSENE EN RARE ee va GEO
. Sterigma strongly eclectic ne a Neenee fal and wide tubular
structure rising collar-like around the ostium. Colliculum absent. Signum small.
Subgenus Spinobactra . . . 5 4 AO
Sterigma, when sclerotized, not Remain € a alle He studie around the
ostium. The ventral rim of ostium usually small or there is a colliculum. Sig-
num larger, basket-shaped . . . . . JR RHM 08
. Sterigma strongly sclerotized, sels stenting € over Fe whole genital
Segment. TLargesspeeies 2 Ne D II, ORARIO.
Sclerotizations not so extensive. . . SR
. Sterigma with a rounded median is Sii in an rommel
COÏICUIUMNE 2: ee GD
Sterigma mesially not lobate. Cle Drm el | A Abe TO
. Colliculum shorter, gradually narrowed downward, its upper edge not extended
laterally; anapophyses longer . . . sola RDD
Colliculum longer, constricted below a aile Sai is ont extended
laterally and narrowed again from above middle to above end; anapophyses
Shorter ode a LRT
. Sterigma, a models Sironi or i leasen “bre plate. Colliculum,
aslonenistronelyssclerotized tube ra nr TIETEN
Sterigma and colliculum not thus. . . . A MOTS
. Sterigma pentagonal, colliculum appearing to sail from its centre, not
distinctly dilated downward. . . . . oe ea ODI
Sterigma transversely oblong, colliculum pri inden from its lower edge,
dilated downwards . . . . De AO
Colliculum well-developed, more or Tess doi maga à to ostium . 14
Colliculum either small, not distincly adjacent to the ostium but beginning
below it, or colliculum ill-defined; if well-defined, then ie and little
selerotizede en a)
Ostium, a es pear- ered stilte asian Callie a be and thick
tube . SEEN AA OR SRR RSL D CEILS
— On; not so lease > all. SU LR AAT
. Colliculum a completely sclerotized diversely shpat fuso SIC RON Fone
Colliculum more sclerotized laterally than in the middle. . . . . . 18
Colliculumispindle:sha pediivery#laroe u 2 Er E
Colliculum tubular. . . V7
. Colliculum rather long, dilated o middle a i glia a transverse
fold along edge of genital segment. . . . Met Ld
Colliculum shorter, gradually narrowed. No transverse fold . . rhabdonoma
A. DIAKONOFF : African species of Bactra 291
18. Folds and structures of lamella antevaginalis less extended, outer as well as
inner usually less than semicircular, the circle being widely open above .
venosana
— hese alde more endet inner folds Lod nine ari concentric
circles; outer structure extended, strongly corrugated, forming more than a
SEMIGITE CM 0. . . punctistrigana
19. Ostium and colliculum little sletten callen upper (anterior) edge
strongly convex, with a median excision; this structure flanked by semicircular
folds and two oval plates of the 7th segment. . . . . . . . endea
— Ostium and colliculum not thus. . . . 1 20
20. Colliculum moderately sclerotized, schamel food br two pata ostium
flanked by an oval structure with double rim and partly sclerotized inner
portion. Lobus analis normal. . . . . dla eo AD
— Colliculum weak, tubular, little sclerotized. Gein scone) projecting for-
wards (ventrad) which is visible only in profile. Ostium flanked by moderate,
strongly sclerotized folds with a hyaline rim. Lobus analis very slender and
elongate ir 3 Mineo cem ar Se7
21. A sclerotized sinuate transverse Lao below aim pe . . … sardonta
NON SU CADA A “ie other species or African Nannobactra
Subgenus Spinobactra nov.
Figs. 2—4
With the superficial characters of Bactra Stephens, but with the valva shaped
differently. Sacculus little indicated, only moderately broader than the base of
cucullus, not swollen, from beyond base with an extremely dense row of long
spines along outer edge; surface of sacculus entirely occupied by punctulations with
fine hairs, although less dense than in other subgenera. Cucullus gradually
narrowed, entirely beset with long spines, especially dense along the base.
Aedeagus broad, curved, moderately long. Cornuti, a small group of spines.
Female genitalia with sterigma strongly sclerotized and forming a complicated
rigid collar-like structure around ostium. Colliculum proper, absent. Signum, a
small denticulate sclerite (similar to that in the subgenus Bactra).
Type-species, Bactra (Spinobactra) spinosa spec. nov.
An interesting novelty of the Bactra group. Apparently a specialized endemic
form. Bactra (B.) pythonia is a transitional form, uniting Spznobactra with Bactra.
The slender insect shows distinct markings that are of a for Bactra unusual
pattern and intensity. It is rather similar to that in pythonia which also suggests
the proximity of these two species.
Bactra (Spinobactra) spinosa spec. nov.
Figs. 2—4
& 10.5—15 mm (holotype 12 mm). Head, antenna, and palpus white, tuft on
vertex tinged creamy, palpus very slightly dusted with greyish laterally. Thorax
white, shoulder mixed with grey. Abdomen whitish-ochreous.
Fore wing oblong-oval, costa rather curved at extremities, less curved in middle,
292 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
apex moderately rounded, termen hardly convex, almost straight, oblique. White,
partly suffused with pale fuscous-grey with a leaden gloss; markings darker fuscous,
partially suffused with tawny. Basal patch to 1/3, indicated by its dark fuscous
edge, ill-defined above, rather obtusely prominent in middle, narrow, slightly sinuate
and vertical below; this patch moderately strigulated with grey-fuscous, hardly
darker than the rest of wing; the prominence of edge of patch forming the first
discal spot, dark fuscous; less than anterior half of costa with very short transverse
strigulae, also obscured by fuscous-grey suffusion; posterior half of costa with 5
rather thick wedge-shaped oblique transverse streaks, separated by the usual pairs
of whitish strigulae; these streaks short, except the fourth which is straight and
traversing wing to before termen below apex; transverse fascia reduced to following
dark fuscous suffused markings, mixed with tawny: a longitudinal moderate streak
along posterior third of upper edge of cell and base of vein 9; an oblong con-
spicuous second discal spot, rounded-prominent below posteriorly and with a
slender upcurved tooth-like appendix, continued across wing obliquely upwards
and branching into fine dark lines along veins; above mentioned longitudinal
streak and second discal spot interconnected by tawny suffusion along closing vein;
ultimate costal streak followed by a slender short line, inwards-oblique and con-
verging with streak below costa; an irregular zigzag blackish line from costa before
apex, to middle of termen, followed by a semicircular outwards-convex line before
lower half of termen; apex beyond black line grey; a conspicuous white area
beyond cell, limited by the described remains of transverse fascia, so as to form
an oblique, suboval well-defined spot with an extension towards costa before apex;
spot in centre of ocellar area present but divided in blackish longitudinal strigulae.
Cilia pale fuscous, dusted with dark fuscous.
Hind wing greyish-fuscous, with a slight bronze gloss, paler towards base, darker
towards apex. Cilia pale fuscous with a grey-fuscous subbasal band.
The long series of males available is variable to some extent, but the species is
characteristically marked and may be confounded only with pythonta. The markings
usually are contrasting, dark on a white ground. Most characteristic is the large
second discal spot, either not incorporated in a transverse fascia, deeply concave
posteriorly and so embracing the white spot beyond cell. Often the edge of the
basal patch is complete, oblique above middle, once more angulate but concave
above dorsum; also the transverse fascia may be complete, its upper portion mode-
rate, oblique, to middle of longitudinal fasciate portion (as described in the holo-
type), second discal spot strongly extended so as to fill posterior part of cell, its
posterior edge concave, containing the white area; usually also the ocellar dot is
dark, conspicuous, and oval. Seldom the wing is infuscated and the markings
obscured.
‘9 12—16 mm (allotype 15 mm). Fore wing slightly broader and more pointed
than in the male. Much paler, the markings being light to pale ochreous-tawny,
otherwise similar to male, but with the pale area beyond cell smaller, more long-
itudinally extended, fasciate and less clearly defined, the entire wing dusted with
pale purple; streaks along veins posteriorly dark and distinct.
Hind wing light greyish-fuscous, slightly tinged brownish.
Other females rather variable (most rather worn); they all are considerably
A. DIAKONOFF : African species of Bactra 293
(7 costa
= processus
LZ =< NC basalis
bristles
N SE = oo A
KEE OMS PIF F N nen
cucullus STA ne
2 DOO 2
aoe DI 9,2 °, ,
Ms [Ue pire
i a ee
terminal thy © aa 2g
cucullus patch He SA
. oo
5 oe
QE \
antibasal Ne |
hairs >
1 a
sacculus
SPC, punctulate
area
Fig. 1. Valva of Bactra (Bactra) species (lacteana), explaining the used terminology of the
parts. Fig. 2. Bactra (Spinobactra) spinosa subgen. et sp. n., holotype, ¢, genitalia
294 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
paler, with markings delicate and rather fainter than in the males. Characteristic
is the small and slender semicircular strigula before lower part of termen, concave
posteriorly (as described in male); the ocellar dot is mostly distinct. One specimen
has the second discal spot suffused with orange-ochreous.
Male genitalia. Tegumen and vinculum rounded, together forming an oval,
vinculum rather broad, band-like. Uncus slender and rather long, corona long.
Socius large. Valva rather broad, cucullus moderate and narrowed, entirely beset
with long spines. Sacculus as described above, entirely covered with sparse punctu-
lations, its outer edge heavily spinose. Aedeagus as described above.
Female genitalia as described above. Lamella antevaginalis strongly corrugated
and of a peculiar shape. Colliculum not sclerotized. Signum very small.
Material examined. Males. South Africa, Transvaal, Pretoria North, 14.11.17
(C. J. SWIERSTRA), 1 4, holotype, gen. no. 4420. Paratypes: Transvaal, Pretoria,
2.11.1911 (D. BURGER), 2 4, gen. 4298, 4412; 31.XII.1909 (4446), 18.111.1910
(4445) (A. J. T. JANSE); Pretoria North, 5.11.1918 (4419), 13.111.1917 (4426),
4.11.1918 (4433), 17.11.1917 (4434) (C. J. SWIERSTRA). Nelspruit, XII.1917
(4421), 11.1918, 2 4, 4422, 4424, 4428 (H. G. BREIJER), Nylstroom, 20.XII.-
1925 (4455, 4468) (A. J. T. JANSE). Three Sisters (near Barberton), 2.III.1911
(4440) (A. J. T. JANSE). Natal: Nkwaleni, 10.1.1916 (4460—62) (A. J. T.
JANSE).
Females. Transvaal, Pretoria. 4.1.1910 (A. J. T. JANSE), allotype, gen. 4318.
Paratypes: the same locality, 23.11.1909 (2873), 5.11.1913 (4435), 1.111.1910
(4431). Pretoria North, 27.1.1917 (4425), 10.11.1918 (4427) (C. J. SWIERSTRA).
Barberton, 20.XII.1910 (4442). Nelspruit, XI.1917 (no abdomen), II.1918
(4439). Natal, Nkwaleni, 10.1.1916, 3.9 (3784, 4310, 4314), 12.1.1916 (4225)
(A. J. T. JANSE).
Subgenus Bactra Stephens, 1834
Bactra Stephens, 1834, Ill. Brit. Ent. Haust., vol. 4, p. 124. — Diakonoff, 1956, Zool.
Verh., no. 29, p. 5 (subgenus).
Aphelia Stephens, 1829 nec Hübner, 1826 (preocc.), Syst. Cat. Brit. Ins., vol. 2, p. 180.
Type-species, Tortrix lanceolana (Hubner, 1796).
This group, presenting so many difficulties with discrimination of its species
in the Palaearctic region (cf. DIAKONOFF, 1962), appears to give even more
trouble in Africa. The northern African representatives of the subgenus are
Palaearctic; they introduce their own, already familiar, problems. The veritable
Aethiopian element, the three species scrupulosa, tradens, and confusa, discussed
below, present difficulties very similar to those of the Palaearctic species, although
to a lesser degree. Also their females are not all too clearly defined. As an exception
to the rule in Bactra, size, shape of the fore wing, colouring and markings of
these species help identification considerably.
Still it may be pointed out that I separated scrupulosa and confusa with the help
of the genitalia first, and was surprised to notice the above mentioned superficial
differences afterwards, now that they were more evident after this sorting out.
A second step was in the opposite direction: I found two different types of males
under confusa first, and after rechecking the slides, discovered the specific differen-
ces of the genitalia afterwards. So three groups of males were separated; subsequent
A. DIAKONOFF : African species of Bactra 295
Fig. 3. Bactra (Spinobactra) spinosa subgen. et sp.n., 9 allotype, bursa. Fig. 4. The same,
sterigma. Fig. 5. Bactra (B.) hebetata Meyr. (= pythonia Meyr.), & holotype, genitalia
296 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
selection of sex partners for scrupulosa and confusa presented now no excessive
difficulties, for these females superficially are similar to the males. Much more
difficult were females of tradens, being intermediate in character. Rearing exper-
iments in some remote future might prove the correctness of my choice.
For the present I prefer to regard the three groups as good species, this in
harmony with my experiences with the Palaearctic species of the subgenus Bactra,
already mentioned in the introduction to the present paper, when referring to
differences in the taxonomic “behaviour” of the three subgenera of the genus
Bactra.
The groups in question — scrupulosa, tradens, and confusa — cannot be sub-
species, because they occur together. I am greatly reluctant to regard them as
“forms’’ or ’’varieties’, because, as far as I am familiar with the Tortricidae, the
differences of the male genitalia in these three species, though not striking, certainly
exceed the limits of differences between infra-subspecific categories.
Bactra (Bactra) pythonia Meyrick, 1909
Figs. 5—8
Bactra pythonia Meyrick, 1909, Ann. Transv. Mus., vol. 2, p. 9, t. 3 fig. 7 (9). —
Clarke, 1953, Meyrick’s Types, vol. 1, p. 267. — Anonymous, 1958, List Types S. & E.
Afr. Museums, vol. 1, p. 57 (holotype” indicated).
Bactra hebetata Meyrick, 1911, Ann. Trans. Mus., vol. 2, p. 227 (8). — Clarke, 1953,
Meyrick’s Types, vol. 1, p. 154. Syn. nov.
Distribution. South Africa: Pretoria; Woodbush Village.
The original description of pythonia is based upon two female syntypes; no
original designation of a holotype had taken place. Where in the “List of Types
in the South and East African Museums” a “holotype” (abbreviated as "H.T.”)
is indicated, actually a lectotype is selected.
This is the first syntype; it was returned to the Transvaal Museum. The second
syntype (a female without the abdomen, but conspecific beyond doubt) was
retained by MEYRICK and is now in the British Museum.
After a critical comparison of a nice series of specimens of the two sexes I am
satisfied that pythonia is the female and hebetata the male of the same species.
The markings, facies and also many localities and dates (cf. below) correspond
closely. The slight superficial differences of the two types fall within the individual
variability, as well as within the sexual dimorphism.
This is a remarkable member of the subgenus Bactra, with markings rather
differing from the patterns which are usual in the Palaearctic species, as summed
up by me lately (1962, p. 5). JANSE’s figure (7 MEYRICK, 1909) is accurate but
the colour is slightly too green. The genitalia are considerably different from those
in the familiar Palaearctic species.
The types may be redescribed as follows.
Bactra pythonia Meyrick (Figs. 6—8).
9 lectotype, 13 mm. Head whitish, edges of vertex suffused with pale tawny-
fuscous. Palpus expanded anteriorly, edge roughish, terminal segment small,
slender, almost concealed; sordid pale ochreous-tawny. Thorax ochreous-fuscous,
tegulae scattered with fine tawny specks. Abdomen fuscous, venter pale fuscous.
A. DIAKONOFF : African species of Bactra 297
Fore wing sublanceolate, rather narrow, costa gently curved, apex moderately
pointed, termen gently concave above, oblique. Ground colour of anterior half of
wing pale grey, of posterior, white. Markings brownish and olive-tawny. Anterior
2/5 of wing whitish-grey slightly dusted with olive and tawny, with about seven
curved and interrupted, rather incomplete transverse brownish lines; every other
line originating from a dark brown triangular dot; this area containing basal patch
which is indicated by deeper and denser olive-tawny suffusion; edge of patch well-
defined, before 1/3, angulate in middle, convex above, concave below angulation;
extreme edge of basal patch suffused with dark fuscous; posterior half of costa
with five dark brown oblique wedge-shaped marks, alternating with minute, less
oblique strigulae: first of these marks on transverse fascia, more distant from
other, fourth continued across wing by a very oblique olive-tawny line; transverse
fascia median, little oblique, formed by a rather irregular broad suffusion, dilated
downward, with moderately suffused edges; fascia narrow and brown on costa,
abruptly dilated and becoming olive-tawny below costa, with a pointed projection
on each side above cell; anterior edge of fascia with a crescentic dark brown pro-
jection on a lower angle of cell, edging white discal dot from below; fascia below
limited by fold; clear white ground colour of posterior half of wing forming the
discal white dot and filling out disc beyond closing vein, thence extended apicad
as a suffused streak directed towards costa before apex; apex with a tawny spot,
centred with black; termen suffused with tawny and dusted with purplish; purple
lines running between and parallel to veins, slightly thickened posteriorly; ocellus
(distinct in left wing) centred with a subquadrate brown mark, indicated by an
irregularly oval brown line along termen and between veins 4 and 5 above mark,
anteriorly curved downwards and running in transverse fascia well below cell;
dorsal margin with a series of dark brown strigulae and dots. Cilia pale grey, dusted
with purple, with a white basal line and purple tips.
Hind wing pale grey, strewn with light tawny-fuscous except towards base,
darker towards apex. Cilia concolorous, infuscated opposite apex.
Female genitalia. Eighth segment not sclerotized, ventral posterior edge with
numerous light wrinkles. Lamella postvaginalis shaped as a slender longitudinal
sclerite, narrowed downward, surface with many small folds; lower extremity of
this sclerite deeply excised. Ostium bursae transversely oval or almost circular,
simple, flanked by two oval refracting and transparent plates, each with a
characteristic bent, also refracting and transparent small thickening of the edge
directed towards ostium. Colliculum present, tubular, rather short, little sclerotized.
Signum absent.
The lectotype is labelled thus: a printed label ’’Pretoria, A. J. T. JANSE”, in the
middle written "26.12.07. 11”. Second label in red ink: “Bactra pythonia M., Type
No. 1001”.
The females are subject to some variation, the basal patch and the transverse
fascia sometimes being obliterate; sometimes there is a black irregular spot in disc
at the place of the obliterated basal patch. In some specimens there is a line along
termen edging ocellus posteriorly, and originating out of a small black streak along
posterior part of vein 5. The ocellus may be dissolved in two horizontal strigulae.
The hind wing sometimes is considerably infuscated.
298 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
59 7 Li : in î fa
FA 29 2 jaf II
N NIN
Fig. 6. Bactra (B.) pythonia Meyr., ® holotype, sterigma. Fig. 7. The same, @, gen.
no. 3557. Fig. 8. The same, 4 , gen. no. 4415
Finally two specimens are melanistic, with the fore wing entirely and evenly
dark fuscous (gen. nos. 4353, 4359).
Bactra hebetata Meyrick (Fig. 5).
& lectotype, hereby selected, 13 mm. Head sordid ochreous-white, sides of
vertex infuscated. Palpus rather slender and smooth, subobtuse, pale grey-fuscous.
Thorax dark fuscous with three pairs of large oval ochreous-whitish spots, more
or less confluent.
Fore wing oblong, gently dilated, slightly broader than in female, costa gently
curved, apex subobtuse, termen faintly convex. Whitish, partly suffused with
A. DIAKONOFF : African species of Bactra 299
grey (tips of scales grey). Markings dark fuscous, slightly suffused with tawny.
Basal patch shaped as in female, but darker fuscous; transverse fascia also fuscous
and darker than in female, shape similar, but tooth of posterior edge above cell
absent (possibly rubbed) and transverse fascia reaching dorsum; costal markings
similar but more extended, also lines originating from them; a round dot in apex,
larger, black; posterior portion of wing more suffused with dark fuscous, thus
rather limiting the white spots described in female, but not quite obscuring them;
distinct are: the white transverse patch beyond closing vein and a stripe from
there towards costa before apex; veins beyond cell, however, suffused with dark
grey, interneural lines extended and black; terminal line distinct throughout;
ocellus as in female but anterior part of bordering line obscured, dark central dot
smaller, confluent with posterior edge of transverse fascia. Cilia light grey with a
white basal line and black submedian and apical lines.
Hind wing rather dark fuscous with a brown gloss. Cilia dark fuscous with a
pale basal line.
Male genitalia. Tegumen rather high, comparable with that of graminivora.
Uncus moderately long. Socius rather large. Valva elongate-oval, with almost
parallel edges, top gradually rounded; hairy and bristly; sacculus swollen, along
the outer edge with rather numerous strong spines, less distinct anteriorly and
posteriorly, punctulate area large, extending over almost the whole surface of the
saccular dilatation, anterior half (left valva) or the whole area (right valva) beset
with strong and long, very dense and numerous spines, directed mesiad and partly
projecting over the cavity of sacculus. The Mc series not distinct, obscured by
numerous long bristles and short spines, densely scattered in this region. The same
concerns Ms series. (The mount no. 4542 is overstained, the dark colouring
obscuring the structure).
The lectotype, present in the Transvaal Museum Collection, has a damaged right
wing and is rather darkly suffused. It is labelled thus. Printed label with green
edge: ”Woodb. Vill., C. J. SwIERSTRA”, in the middle written: 15.12.1909”. A
written label "G(enitalia) 4542”.
Usually a clearly marked species, with distinct white spot beyond cell and white
streak towards end of costa, contrasting with deep tawny or brown of the transverse
fascia and its crescentic projection; the ocellus is also characteristic.
Material studied. South Africa, Transvaal: Pretoria, Pretoria North, N.E.
Pretoria, Heidelberg, Naboomspruit, Woodbush Village, Zoutpansberg, Waterval-
Onder, Nelspruit, Kranspoort. Natal: New Hanover, Karkloof. Cape
Province: Durban, Congella near Durban. Southern Rhodesia: Bulawayo.
(A. J. T. JANSE, C. J. SWIERSTRA, H. G. BREIJER, LEIGH). 25 4,13 9 (Genital
nos. except types: 3 3521, 4411, 4413 —4418, 4423, 4430—32, 4438— 39, 4444,
4449, 4451-54, 4458, 4469. 9 2881, 3558, 3649, 4347, 4353, 4359, 4376,
4429, 4435 — 36, 4441, 4443, 444748, 4464, 4467, 4470, 4545).
A widely distributed species. Its position is very interesting for it is intermediate
between the subgenera Spinobactra and Bactra, but closer to the latter and therefore
incorporated in Bactra. Still it does not fit in the older conception of the subgenus,
for the female genitalia and also the sacculus of the male are peculiar and very
unusual.
300 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
Bactra (Bactra) dolia spec. nov.
Figs. 9—11
‚9 20—21 mm (holotype 21 mm). Head, palpus and thorax pale ochreous,
palpus strongly dilated, with a roughish edge, slightly mixed with tawny. Abdo-
men pale ochreous.
Fore wing oblong, rather narrow, costa gently curved towards extremities, apex
pointed, termen hardly sinuate, strongly oblique. Pale ochreous, dusted with light
ochreous-tawny, markings light and dark fuscous. Costa with numerous light
tawny strigulae, oblique and very slender, reaching to cell; dorsum and termen
more dusted with light tawny, terminal area denser; a dark fuscous little suffused
median stripe along lower half of cell, from base to closing vein; a fuscous-tawny,
much lighter suffusion filling out remainder of cell and extending beyond cell so
as to reach below apex; an interrupted narrow brownish line; a faint blackish dot
in apex; a series of dark fuscous dots along dorsum. Cilia light tawny with a sub-
basal and a supramedian brownish line.
Hind wing with a strong golden gloss, suffused with light purplish-fuscous,
dorsum paler. Cilia pale golden.
All specimens are slightly rubbed and very uniform. Only the Transvaal Museum
specimens have paler hind wings and a more contrasting median dark streak; the
specimen from St. Lucia Lake has a conspicuous black dot in apex, continued as
a narrow bar over cilia.
Female genitalia resembling those of scrupzlosa but differing as follows.
Anapophyses and postapophyses extremely long. Lobi anales longer and more
slender along the proximal portion. Ostium bursae very similar to that in scrupu-
losa, but on top of a much larger and considerably sclerotized process or column;
these sclerotizations conspicuous, forming an L-shaped sclerite on each side, lower
half extended into an oval plate. Lamella postvaginalis forming a strong cap above
ostium, above this cap the wall is strongly punctulate and with 1—5 articulating
bristles on each side (never present in scrupulosa). Signum absent.
Discovery of the corresponding male will be interesting.
Material studied. In the British Museum. South Africa, Natal: Camperdorn,
4.1908 (L.), holotype, @, gen. no. 3468, the same III.1908 (L.), paratype 9,
gen. no. 3614 (MEYRICK Collection).
In the Transvaal Museum. Natal, Umkomaas, 1.1.1914 (A. J. T. JANSE), 1 9,
paratype, gen. no. 4301. St. Lucia Lake, X.1917, 1 9, paratype, gen. no. 4409.
49.
Bactra (Bactra) scrupulosa Meyrick, 1911
Figs. 12—15
Bactra scrupulosa Meyrick, 1911, Ann. Transv. Mus., vol. 2, p. 227. — Clarke, 1955,
Meyrick's Types, vol. 1, p. 281. — Anonymous, 1958, List Types S. & E. Afr. Museums,
vol. 1, p. 57 (’’holotype’’ and “paratype” indicated).
Distribution. South Africa: Woodbush Village, Albert Mine.
4 lectotype, 13 mm. Head whitish, vertex pale ochreous with a white median
stripe. Palpus pale ochreous, strongly dilated, upper edge and apex roughish; pale
A. DIAKONOFF : African species of Bactra 301
Fig. 9. Bactra (B.) dolia sp.n., © holotype. Fig. 10. The same, 9 paratype, gen. no. 4409.
Fig. 11. The same, sterigma, paratype, gen. no. 4301. Fig. 12. B. (B.) scrupulosa Meyr.,
8 lectotype, gen. no. 4186
302 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
ochreous, terminal segment slender, short, blackish. Antenna pale ochreous. Thorax
pale ochreous, somewhat streaked longitudinally with white.
Fore wing narrow, sublanceolate, costa gently curved throughout, more curved
anteriorly, apex moderately pointed, termen hardly concave, oblique. Ground
colour glossy white, marbled, and over terminal third, suffused with pale tawny-
ochreous. Costa with a series of large and rather irregular marks, some 12 of them,
dark fuscous along anterior half, fuscous mixed with tawny along posterior half;
basal patch indicated by an inwards-oblique tooth-shaped mark above fold beyond
base of wing and by an irregular obliquely-subquadrate dark fuscous spot just
before 1/4, situated in middle of disc, upper edge concave, outer and lower edges
rounded; transverse fascia represented by a moderate tawny-fuscous, oblique and
zigzag mark from below costa before middle, to below lower angle of cell, rather
well-defined, in left wing narrowly connected with costa; this mark followed by a
small and short horizontal streak halfway between cell and tornus; wing beyond
cell suffused with pale tawny-ochreous, with a faint indication of a regularly oval
deeper tawny streak from lower angle of cell to vein 7 beyond its base, along 7
to before termen, thence curved down and running to termen above tornus;
blackish dusting forming fine lines above veins 7 and 6; a round tawny spot in
apex; dorsal edge with small irregular fuscous blotches. Cilia greyish with a sub-
basal and a postmedian fuscous lines.
Hind wing pale grey, from middle towards apex becoming rather dark grey.
Cilia glossy white, with an interrupted pale fuscous subbasal line.
Lectotype, labelled with a printed, green-edged label: "Woodb. Vill., C. J.
SWIERSTRA”, in middle written: ”11-12.1909”’. A label in red ink: ’’Bactra
scrupulosa Meyr., Type no. 2623.” Genitalia no. 4186 4. ’Cotype” a ¢ with a
quadrate label in red ink: Bactra scrupulosa Meyr. Cotype No. 1000” label in
black ink: "Alb. mine 9.11.07 11” (gen. no. 4189). This specimen is much darker,
strongly marbled and suffused with bright tawny partly dusted with dark fuscous,
markings suffused; basal patch complete, edge strongly angulate, to 1/3, upper
half more oblique than lower half; transverse fascia shaped as an oblique straight
band, from costa before middle to below end of cell, becoming darker brown
downwards, merging in an indistinct pale tawny-ochreous marbling along dorsal
third of wing; apical streak broad in middle; anterior end connected by a tawny
line with 2/3 of costa; two similar costal lines, parallel, between basal patch and
transverse fascia; costa slightly sprinkled with black, several vertical series of black
strigulae before termen below apex. Cilia pale tawny along termen, towards apex
with a distinct white basal streak.
Hind wing similarly obscured but more fuscous-tinged.
The species usually is rather mottly marked and then may be referred to the
’’maculate” type (DIAKONOFF, 1962, p. 7, t. I, fig. 1); with a well-defined basal
patch and a transverse fascia including the second discal spot; this latter, however,
is not V-shaped but simply forms a straight and oblique continuation of the
transverse fascia. An apical streak is not developed.
The long series available is variable; it is chiefly characterized by the small size
and the narrow pointed wings of the two sexes. Furthermore, all males which are
not too much rubbed show a transverse fascia originating from middle of costa,
A. DIAKONOFF : African species of Bactra 303
usually more distinct and thicker than the parallel costal strigulae; these strigulae
are moderate in number, not very slender and rather distant; the fascia runs
obliquely across the wing and ends in a rather broad stripe across the lower angle
of cell, including the second discal spot. The connection of this spot with the
costa may be very narrow, but it is almost always traceable in not rubbed specimens.
The basal patch in male is mostly distinct, its edge well-defined, at least along the
upper half and is but seldom reduced to a transverse fasciate dark first discal spot.
The hind wing in the males is varying from dark fuscous (seldom) to pale
greyish-fuscous, with a whitish base.
Both the fore and the hind wings are narrower and more pointed than in
confusa.
The females are slightly more variable than the males, but have also narrower
and longer wings than in confusa. They are usually more differing from this
species than the males.
The following is a description of a characteristic female.
18 mm, neallotype. Head pale ochreous suffused with tawny, except on face.
Palpus rather long, strongly dilated, edge roughish, pale ochreous, suffused with
tawny. Antenna dark brown. Thorax pale ochreous, slightly spotted with pale
tawny, tegula light; metathorax white. Abdomen pale ochreous.
Fore wing elongate, narrow, costa considerably sinuate, apex slightly produced,
pointed, termen sinuate, oblique. Pale ochreous, along costa strigulated, elsewhere
dusted or suffused, with light tawny. Costal strigulae slender, becoming longer
along posterior half of costa and deeper tawny towards apex; a median longitudinal
streak from base to apex, of bright tawny suffusion, dusted with minute black
dots (tips of scales), this streak dilated beyond base, not quite filling cell, beyond
cell slightly narrowed; termen suffused with paler tawny; apex with a blackish
point, termen with a faint brown line, a series of blackish dots along dorsal edge;
apex with a blackish dot. Cilia in tornus pale ochreous; elsewhere cilia tawny
dusted with brownish and with white basal line, followed by a postmedian and
a subapical narrow whitish lines.
Hind wing whitish golden, posterior half deeper golden tinged and with a
faintest fuscous suffusion. Cilia whitish golden, paler towards dorsum, more
ochreous-tinged along termen and around apex, a shadow of a subapical fuscous
line; opposite apex cilia with brownish subbasal and subapical short lines, apex
with a few dark dots.
Female specimen redescribed: Pretoria, 8.1.1910, (A. J. T. JANSE), gen. no.
4373.
Besides the described female which is of the fasciate type, which is the most
common, there is also the reduced type present, derivable from the maculate type as
described for the male, but with markings for the greater part obliterate; there are
also a few pale females, the unicolorous, or almost unicolorous type.
Female genitalia are very close to those of the Palaearctic Bactra lacteana Caradja,
but differ by the invariable absence of a signum. The ostium bursae is perhaps a
trifle wider than in /acteana, the colliculum part is very slender, the edge of the
ostium usually has several small transverse folds. The lamella postvaginalis some-
times is moderately but distinctly sclerotized around the lateral corners of the
304 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
Fig. 13. Bactra (B.) scrupulosa Meyr., & syntype, gen. no. 4189. Fig. 14. The same, @
gen. no. 4373. Fig. 15. The same, bursa
ostium; the ’’cap’’ above the ostium is as in lacteana, convex above, with several
longitudinal folds, crowned by some distinct punctulation of the wall, sometimes
shaped as small triangles, ears’, at the sides, exactly similar to those in lacteana.
The lamella antevaginalis may be ill-defined or slightly indicated by a faintest
sclerotization and then shaped as a subtriangular small plate with or without a
A. DIAKONOFF : African species of Bactra 305
small curved fold on each side of the colliculum part of the ductus bursae; lateral
ends of the ostium cap are then continued as the sides of that triangle. The
apophyses are moderately long and slender. Actually the ostium is situated on top
of a moderate prominence, visible only in profile.
Material studied. Besides the lectotype and the original syntype, mentioned
above, the followings males. In the British Museum. South Africa, Transvaal:
Pretoria. Natal: Frere, 3600 ft.; Weenen, Kimbolton Estcourt, 13.XI untill 22.11
(A. J. T. JANSE, C. J. SWIERSTRA, MARSHALL and HUTCHINSON, in MEYRICK and
WALSINGHAM Collections).
In the Transvaal Museum. Transvaal: Pretoria, Fountain Grove, New Smitsdorp,
Woodbush Village, Elandshoek, Modderpoort. Natal: Karkloof, Rietvlei,
Eshowe, Cathedral Peak. Cape Province: Stellenbosch, Alexandria (Langebos)
DIN VER (Collected ss by A/R [ANSE Gi CHJ SWIERSTRA, MOLTO MNS:
VILJOEN, Ch. K. BRAIN, A. CAPENER, L. VARI). Genit. nos: ¢ 2872, 3461, 3518,
3559, 3637—38, 3642—43, 364748, 3777, 3782—83, 4217—18, 4220—22,
4224, 4226, 4229—30, 4256, 4259, 4261, 4265, 4271, 4338, 4340, 4368, 4371,
4408, 4465, and T.M. 4541.
Females. In the British Museum. Transvaal: Pretoria, Barberton. Natal: Weenen,
Kimbolton Estcourt. I—II (A. J. T. JANSE, H. THOMASSET, HUTCHINSON, in
MEYRICK and WALSINGHAM Collections).
In the Transvaal Museum. Transvaal: Pretoria, Barberton, Pretoria North,
Nelspruit, Waterval-Onder, Nylstroom. Natal: Karkloof, Rietvlei. Cape Province:
Kleinmond 25.X—23.III (A. J. T. JANSE, C. J. SwIERSTRA, H. G. BREIJER, H. G.
IDICKSON) Genie Ossi) 2871290534 07352803553 30M:
3795—6, 4223, 4227—8, 4245—6, 4248, 4255, 4257—8, 4260, 4262—3, 4266,
4290, 4317, 4324, 4325, 4328, 4332, 4346, 4356, 4360, 4373. In total 33 &,
DAMON
Bactra (Bactra) tradens spec. nov.
Figs. 17—18
& 13.5—17 mm (holotype 16.5 mm). Head pale ochreous, face creamy. Palpus
creamy, laterally mixed with grey towards dorsal edge which also bears two well-
defined black transverse marks; terminal segment grey. Thorax pale ochreous,
tegula mixed with brownish along edge and on apex. Abdomen ochreous-fuscous.
Fore wing oblong-triangular, distinctly dilated, costa curved, apex pointed,
termen sinuate, long, oblique. Pale ochreous, evenly suffused with pale leaden, all
markings edged with creamy. Markings light tawny and dark brown, of the
’tigroid” type. Costa with some 20 dark brown subtriangular transverse marks,
each other mark continued as a slender tawny very oblique transverse strigula;
along anterior half of costa these strigulae run more or less straight across cell,
those along posterior half run parallel to preceding at origin but below costa turn
horizontal and run towards termen, thence curve down and become almost vertical;
ultimate two or three strigulae run to termen and along this to tornus; posterior
extremity of costa with a conspicuous whitish spot; first discal spot somewhat irre-
gular, dark brown with two teeth; second discal spot elongate-semioval, almost
black; stigma not defined; space before termen somewhat more suffused with
306 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
tawny, appearing darker, space above dorsum less so suffused, lighter; a very faint
ocellar dark dot; a suffused short blackish streak through apex. Cilia dark grey
with a narrow basal, a faint antemedian, and an ill-defined subapical white lines.
Hind wing rather pale fuscous-purplish, becoming paler, almost transparent,
on dorsum. Cilia sordid pale fuscous with a paler base and a grey subbasal streak.
Other males all show the same "tigroid’” pattern with numerous, well-defined
fine strigulae and limited discal spots; one specimen is rather bright tawny tinged,
another rather suffused with darker leaden, a third shows a contrasting median
blackish streak from base to apex, with an obtuse prominence on end of cell
(’tigroid-fasciate”, 4362).
9 16—19 mm (allotype 17.5 mm). Fore wing narrower, less dilated, but apex
more pointed and termen sinuate and more oblique than in male. Colouring more
variable. In the allotype fore wing strongly suffused with bright tawny-ochreous,
finely reticulate and marbled with dark purplish; cell filled out with purplish;
markings reduced to short dark strigulae along costal edge.
Hind wing pale fuscous, becoming paler towards base.
There are four female paratypes with similar genitalia and appearance; they are
somewhat brighter tawny suffused and have a more distinct, although strongly
suffused, median dark stripe. Three more females have less typical genitalia; these
specimens are lighter coloured. It seems advisable to exclude them from the para-
type series.
Male genitalia. Perhaps best comparable with the Palaearctic furfurana, but
distinct. Also close to confusa. Tegumen rather low. Uncus moderate, curved, top
pointed. Vinculum moderate. Valva broad and short. Cucullus bristles normal or
rather weak. Sacculus moderate, somewhat smaller than in confusa, spheroidal,
outer edge of cucullus is continued over the surface of the sacculus, thus forming
a distinct small ridge or fold, the Ms series thus being unusually accentuated; this
small ridge continues almost to the punctulate area. Terminal patch present but not
dense; Mc series is not clearly defined or separated from the fine setae between
this series and the terminal patch, in very much the same way as in confusa. The
punctulate area is limited, shorter, narrower at the top, with 3—4 short curved
spines; the group of basal hairs is clearly extended and entirely united with the
punctulate area. The antibasal group of setae large, setae numerous and long (as
in confusa). Aedeagus and caulis both rather long and sclerotized.
Female genitalia are not easily discriminated from these of confusa or scrupulosa,
this forming the greatest difficulty with the identification of the species of the
present paper. Actually intermediate between confusa and scrupulosa. Ostium
wider than in the latter, less wide than in the former species. The rostral edge of
the eighth tergite clearly sclerotized, uniting the bases of similarly sclerotized
anapophyses as a bow or a semicircle, thus differing from scrupulosa where these
parts are not sclerotized. The lateral sclerites of the ostium are not defined or
indicated by very faint fragments of their caudolateral edges; however, each has a
small peculiar dark and tubular appendix caudomesially, not developed in the
other two species. Ductus and corpus bursae normal. Signum absent.
Material studied. Males. South Africa, Transvaal: Pretoria, 30.1.1910 (A. J. T.
JANSE), holotype, gen. no. 4311. Paratypes: Pretoria, 3 and 29.XI.1909 (4312,
A. DIAKONOFF : African species of Bactra 307
Fig. 16. Bactra (B.) confusa sp.n., &, holotype, genitalia. Fig. 17. B. (B.) tradens sp.n.,
4, holotype, genitalia
308 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
4326); 17.1.1910 (4313, 4323); 17.XI1.1909 (T.M. 4544); 14.X.1917 (4362);
Mahuba’s Kloof, 1050 m, 18.1.1925 (4357) (All collected by A. J. T. JANSE).
Natal, Karkloof, 11.1.1917 (4336) (same collector). Cape Province, Swellendam,
IV.1943 (4336) (C. G. Dickson). (In the Transvaal Museum). Transvaal, Pre-
toria, 2 and 8.XII.1909 (JANSE), 2 paratypes, gen. 3464 and 3465. (In the British
Museum).
Females. Transvaal: Waterval-Onder, 11-13.X.1951 (A. J. T. JANSE), allotype,
gen. no. 4404. Paratypes: Transvaal: Nelspruit, XI.1917 and 11.1918 (H. G.
BREIJER), gen. 4246, 4247 and 4248; Pretoria North, 23.111.1927 (C. J. SWIER-
STRA), gen. 4245. Pretoria, 3.11.1910 (A. J. T. JANSE). Besides, the following
females: Pretoria, 3.X.1907 (4333), 8.11910 (4373); Nylstroom, 4-5.111.1954
(4407). (All collected by A. J. T. JANSE). In total 12 & and 8 9.
The species is intermediate between scrupulosa and confusa and also closely
allied with furfurana. From the former two species it may be separated by the
genitalia, from the latter, by entirely different facies and markings of the two
sexes and also by the different sacculus of the male and the absence of a signum
in the female.
Bactra (Bactra) confusa spec. nov.
Figs. 16, 19
4 12-15 mm (holotype 14 mm). Head, palpus and thorax pale ochreous-
fuscous, tinged olive, face whitish, palpus laterally slightly dusted with grey.
Abdomen pale fuscous.
Fore wing oblong, subovate, broad, costa rather curved throughout, less curved
in median portion, apex pointed, termen long, straight, oblique. Whitish-ochreous,
generally with olive tinge, markings being formed by fine olive reticulation, only
partly mixed with fuscous. Costal markings not numerous, about 7—8 strigulae,
dark fuscous on costal margin, pale olive thence, moderately oblique, faint along
anterior half of costa, well-defined along posterior, throughout alternating with
minute dark lines; basal patch not reaching to 1/3, indicated by a narrow dark
fuscous edge only, which is angulate slightly above middle, rather obtusely, and
gently sinuate above dorsum; transverse fascia well-traceable, from middle of
costa to just above end of fold, strongly suffused, olive-tinged, mixed with fuscous;
this fascia moderate on costa, including a very irregular and strongly suffused
second discal spot which is obliquely transverse; stigma hardly perceptible and
ill-defined; lower third of posterior edge of fascia abruptly well-defined, serrate
and little oblique; a fuscous suffused streak from apex to end of termen, gradually
attenuated downwards, marked with a few blackish vertical strigulae. Cilia fuscous,
a basal line and some tips of scales whitish.
Hind wing unicolorous dark grey-fuscous, with a very faint golden gloss. Cilia
grey-fuscous with a whitish basal line.
The most conspicuous features of the species are broad and pointed wings with
not sinuate termen, the fuscous-olive marbled markings and the unicolorous dark
hind wings. The series of males is uniform, one specimen is rather suffused with
brighter ochreous-olive and has a pronounced row of dark brown costal dots.
Q 16—18 mm (allotype 17.5). Head and palpus paler, whitish-creamy, woolly
A. DIAKONOFF : African species of Bactra 309
by rather loose scales, palpus finely dusted with grey, terminal segment grey. Fore
wing slightly broader, costa a trifle more curved. Very similar to male but paler,
less suffused, markings therefore more pronounced, thicker, unicolourous pale
tawny-olive, without dark marks along costal margin; strigulae along posterior part
of costa more complete, reaching to termen, one from beyond 1/3 of costa even
traceable to tornus, forming a loop in centre of disc; transverse fascia including
a second discal spot of more usual shape; semicircular, with a traceable stigma;
first discal patch is dissolved so as to form fuscous marbling.
Cilia and hind wing exactly as in male.
Another female (4385) is coloured thus. Rather densely suffused with tawny-
olive, almost entirely obscuring the strigulae, except along costal edge where they
are distinctly dark brown and short; transverse fascia similarly obscured by olive
suffusion, only the second discal spot is distinct, being dusted with dark brown,
as also is the terminal streak in apex and along its lower half; some dark dots
along dorsum.
Other females similar, two are somewhat brighter coloured and with dark
fuscous first discal spot rather extended along fold.
Male genitalia. Rather similar to those of the Palaearctic B. (B.) lacteana Car.,
but distinctly different. Tegumen and socius normal. Uncus rather long (longer
than in /acteana), top distinctly more pointed. Valva similar, rather broad and
short, with a rounded top. Cucullus bristles in a broad row which becomes less
and less dense towards sacculus; terminal patch is present, broad but not dense
and somewhat irregular, continued far up the surface of sacculus; anterior edge
of this patch beset with long and straight spines, then abruptly changing into Ms
series which is seemingly double but sparse, then becoming single; the Mc series
is rather ill-defined, occupying a broad area on and between the ridge of the
processus basalis and extending halfway towards the Ms. The punctulate area
similar to that of lacteana, large, semioval, top gradually rounded. Spc, spines
rather long, slender and straight. The group of basal hairs is moderate but larger
than in /acteana. There is another characteristic feature: a group of hairs on the
opposite side of the sacculus just below the terminal patch, which I indicate with
the name of antibasal hairs. This group consists of rather numerous long and thin
setae. (In lacteana there are only a few very small setae there). The outer surface
of sacculus is strongly wrinkled because of numerous scars of scales. Aedeagus and
caulis rather long.
Female genitalia. Of a type similar to that of scrupulosa, but clearly distinct.
Sterigma forming a prominence, much broader than in that species; this structure
is flanked by two subcrescentic plates, clearly sclerotized, gradually dilated up-
wards, lower portions forming a semioval horizontal plate on each side. Cap of
the ostium (lamella postvaginalis) wide, with several longitudinal folds, forming
a smaller cap on each side, moderately sclerotized. Wall above cap only with a
few wrinkles laterally. Ostium wide, single, calyciform. Ductus bursae narrow.
Signum absent.
Material studied. South Africa, Transvaal: Wylie's Poort, 6.XI.1920 (C. J.
SWIERSTRA), 1 &, holotype, gen. no. 4386. The same locality and collector,
6.XI.1920, 1 9, allotype, gen. no. 4387. All other paratypes: the same locality
310 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
and collector, 3-10.XI.1920, 1 4, 49, nos. 4383 4, 4380—1, 4384—5. Natal:
Sarnia (Mrs. GURRY), XI.1913, 1 4,119 (4383 4, 3786); the same locality.
10.11.1912 (WILLIAMSON), 1 2 (4278) (Coll. JANSE). Umkomaas, 4.XI.1920
(C. J. SWIERSTRA), 1 & (4382); the same, 23.1.1914 (A. J. T. JANSE) 1 &
T.M. 4642). Karkloof: 12-20.1.1917 (A. J. T. JANSE), 2 $ (3797, 4475). Natal,
Pinetown (L.), 1.1909 (Meyrick Coll., British Museum), 1, 9, gen. no. 3460.
Dei LOMO
The present species could be separated without much difficulty. After having
sorted out the specimens with the aid of the genital characters I was struck by
the uniformity of the so selected series of the rather large, broad-winged insects.
Intermediate between tradens and scrupulosa.
Bactra (Bactra) furfurana (Haworth, 1811)
Tortrix furfurana Haworth, 1811, Lep. Brit., p. 466.
Bactra furfurana: Wilkinson, 1859, Brit. Tortr., p. 147. — Zeller, 1875, Verh. zool.-bot.
Ges. Wien, vol. 25, p. 41. — Fernald, 1882, Trans. Am. Ent. Soc., vol. 10, p. 29.
Grapholitha (Aphelia) furfurana: Heinemann, 1863, Schmett. Deutschl., Kleinschm.,
VOL ik, joy 35.
acutana Eversmann, 1844, Faun. Lep. Volgo-Ural, p. 529 (non binom.).
Phoxopteris lamana Zeller, 1846, Isis, vol. 30, p. 257.
Tortrix (Aphelia) scirpana Herrich-Schäffer, 1849, Syst. Bearb. Schmett. Eur., vol. 4,
p. 243.
Tortrix (Aphelia) pauperana Herrich-Schäffer, 1849 (nec Haworth, 1811), Suppl. p. 302.
Sciaphila canuisana Milliére, 1847, Rev. & Magas. Zool., ser. 3, vol. 2, p. 247. — 1875,
Ann. Soc. ent. Cannes, t. 1, f. 6.
Distribution. Northern Africa, Morocco: Buhasen. Palaearctic and Nearctic regions. Must
have been introduced from Europe.
Bactra (Bactra) lanceolana (Hübner, 1796)
Tortrix lancealana Hübner, 1796, Samml. europ. Schmett., vol. 7, Tortr., t. 13, f. 80.
Ancylis lanceolana: Hübner, 1825, Verz. bek. Schmett., p. 376.
Phoxopteris lanceolana: Treitschke, 1830, Schmett. Eur., vol. 8, p. 232, no. 1.
Phoxopterix lanceolana: Duponchel, 1834, Hist. nat. Lép., p. 339, t. 253, fig. 1.
Tortrix (Aphelia) lanceolana: Herrich-Schäffer, 1849, Syst. Bearb. Schmett. Eur., vol. 4, p.
243. — Suppl. 1849, p. 317.
Bactra lanceolana: Wilkinson, 1859, Brit. Tortr., p. 115. — Stainton, 1859, Manual, vol.
2, p. 226. — Fernald, 1882, Trans. Amer. Ent. Soc., vol. 10, p. 28.
Grapholitha (Aphelia) lanceolana: Heinemann, 1863, Schmett. Deutschl., Kleinschm., vol.
1, p. 134.
Tortrix dibeliana Hübner, 1800, Samml. europ. Schmett., vol. 7, Tort, p. 272.
Tortrix pauperana Haworth, 1811, Lep. Brit, p. 469.
Tortrix expallidana Haworth, 1811. Lep. Brit., p. 469.
Tortrix egenana Haworth, 1811, Lep. Brit, p. 469.
Tortrix egestana Haworth, 1811, Lep. Brit., p. 470.
Tortrix lanceana Frölich, 1828, Ent. Tortr. Würt., p. 98.
Bactra nigrovittana Stephens, 1852, List Brit. Anim., vol. 10, p. 99.
Fig. 18. Bactra (B.) tradens sp.n., ® allotype, sterigma and ovipositor. Fig. 19. B. (B.)
confusa sp.n., ® allotype, sterigma and ovipositor. Fig. 20. B. (B.) crithopa Diak., &
holotype, genitalia (fig. 20 after Diakonoff, 1960)
A. DIAKONOFF : African species of Bactra 311
312 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
Bactra (Aphelia) egenana Kennel, 1900, non Haworth 1811, Iris, vol. 13, p. 264.
Bactra (Aphelia) fumosana Kennel, 1900, Iris, vol. 13, p. 263.
Bactra fumosana: Kennel, 1910, Palaearkt. Tortr., p. 472—473, t. 18, fig. 73a. — Rebel,
in Staudinger & Rebel, 1901, Catal. Lep. Pal. Faun., vol. 2, p. 113.
Bactra (Bactra) fumosana: Diakonoff, 1956, Zool. Verh., no. 29, p. 16, fig. 16 (gen. ®
type fig.).
Distribution. Northern Africa, Algeria, Morocco. Palaearctic region. Canada. Obviously
introduced in Northern Africa from Europe.
Bactra (Bactra) graminivora Meyrick, 1922
Bactra graminivora Meyrick, 1922, Exot. Microl., vol. 2, p. 521 (4 2, Bengal). —
Fletcher, 1932, Life histories Ind. Microl., ser. 2, p. 24, t. 14 figs. a-d (biol., food-plants).
— Diakonoff, 1950, Bull. Brit. Mus., Ent., vol. 1, p. 287, t. 6 fig. 24, t. 7 figs. 34, 36
(lectotype select, gen. 4 9 fig.). — Amsel, 1958, Beitr. nat. Forsch. S.W. Deutschl., vol.
17, p. 76 (N. Arabia).
Bactra cyperana Amsel, 1951, Bull. Soc. Sci. Nat. Maroc, vol. 31, p. 68, fig. 4 (8,
Morocco).
Bactra mediterraneana Agenjo, 1952, Faunula Lep. Almeriense, p. 99, t. 4 figs. 32, 33, t.
12 figs. 6, 7 (4 2, Spain, Almeria).
Bactra (Bactra) graminivora: Diakonoff, 1956, Zool. Verh., no. 29, p. 18, fig. 10. —
1959, Bijdr. Dierk., fasc. 29, p. 179, figs. 6, 7 (cyperana, mediterraneana syn.). — 1962.
Zool. Verh., no. 59, figs. le, 18—20, 29, 30; t. III figs. 17—20. — 1963, Ann. Mus.
Wien, vol. 67 (in press) (4, Nubia).
Distribution. Canary Islands, South Spain, Malta, Algeria, Morocco, Egypt, Sudan, Nubia,
Gambia, French Congo, Arabia, Iraq, Mesopotamia, Caucasus, Caspian region, Tadzhikistan,
Bukhara, North Persia, Afghanistan, Hyderabad, Bengal.
This widely distributed Asiatic species apparently is a straggler in Africa. The
incontinuous distribution suggests that it is rare and local.
The species is represented by the maculate form in the males with more or less
pronounced discal markings characteristic for this species, while the females are,
as usual, of the unicolorous form, with one exception, a vittate female, a form
not familiar to me in graminivora so far.
Material studied. Transvaal Museum. Transvaal: Pretoria, 24.X.1915 (A. J. T.
JANSE), 1 &, gen. no. 3785; 1 ©, gen. no. 3415. Natal: Bela Vista, XI.1916
(C. J. SWIERSTRA), 1 2.
Vienna Museum. French Congo: Insongo, 27.111.1927 (WEIDHOLZ), 1 9, 4483.
Sudan, Sungikai 19.111.1914 “Bactra lanceolana Hb., det. Meyrick”, 1 ‚9, 4484;
Debri, 25.111.1914, 1 4, 4485 (as above); Rosti, 21.IV.1914 (do.), 1 4, 4486;
Kororak, 31.111.1914, 2 9, 4487—8 (EBNER). In total 3 4,519.
British Museum. West Africa: Gambia, Bathurst, XI.1884 (CARTER), WAL-
SINGHAM Collection, no. 893, 1 9, gen. no. 3650.
Bactra (Bactra) crithopa Diakonoff, 1957
Fig. 20
Bactra (Bactra) crithopa Diakonoff, 1957, Mem. Inst. Sci. Madagascar, ser. E, vol. 8, p.
272, fig. 29, t. 8 fig. 20 (8).
Distribution. Reunion.
A. DIAKONOFF : African species of Bactra 313
The species of which only the single male is known, is easily recognisable by
the Mc series formed not by hairs but by small spines, and by unusually large
socii.
Subgenus Chiloides Butler, 1881
Chiloides Butler, 1881, Ann. Mag. Nat. Hist., ser. 5, vol. 7, p. 392. — Meyrick, 1885,
Trans. New. Zeal. Inst., vol. 17, p. 142. — Fernald, 1908, Gener. Tortr., p. 44, 58. — Flet-
cher, 1929, Mem. Agr. Ind., Ent., vol. 11, p. 30, 46. — Diakonoff, 1956, Zool. Verh., no.
29, p. 19 (subgen. of Bactra).
Type-species, Chiloides straminea Butler, 1881.
For a short discussion of the subgenus may be referred to the introduction.
Bactra (Chiloides) jansei spec. nov.
Figs. 21—23
8 13—15 (holotype 15 mm). Head creamy. Antenna fuscous. Palpus pale
ochreous. Thorax creamy, tegula infuscated, metathorax silvery-white. Abdomen
pale ochreous, venter whitish.
Fore wing oblong, suboval, costa considerably curved throughout, apex pointed,
termen sinuate, oblique. Creamy, rather densely dusted with purplish-brown, except
an area in disc beyond cell extending halfway towards termen and limited by veins
3 and 8. Markings of the “tigroid” type, slightly reduced, tawny and brown.
Costa with 6—7 thicker oblique strigulae, slightly undulate, some in the median
portion of costa furcate towards wing margin (a very unusual feature; this
furcation must be more or less incidental, as it is not congruent in right and left
wings); these strigulae reach to cell and to vein 8, and alternate with minute
shorter lines; penultimate costal streak forming a curved line across wing to
tornus; basal fourth of wing rather strongly infuscated; first discal spot obliterate,
second discal spot linear, a longitudinal dark fuscous strigula just above lower
angle of cell; veins beyond cell finely streaked with brownish; a well-defined
terminal streak, gradually dilated upwards, merging in a subtriangular and suf-
fused subapical dark brown spot, being a reduced apical streak, including a small
black streak through apex; a creamy spot on costa above apex. Cilia creamy with a
suffused fuscous subbasal line.
Hind wing pale golden-fuscous, becoming very pale towards base, deeper fuscous
towards apex, glossy. Cilia whitish, with a pale fuscous basal half.
The male series is rather variable, with more or less defined strigulation, some
are pale fuscous, other pale ochreous tinged. One specimen (4327) is of the
maculate type, with the fine strigulation noticeable but not conspicuous and with
the both discal spots large, well-defined, fuscous, roundish, second spot larger and
lighter than first. Sometimes (4322) the dorsal half of wing is suffused with
purplish-fuscous. In paler specimens there is a contrasting black dot in apex.
Q 13—17 mm (allotype 17 mm). Head and thorax fuscous mixed with creamy.
Palpus pale ochreous, with an oblique median greyish stripe. Abdomen ochreous-
fuscous.
314 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
A. DIAKONOFF : African species of Bactra 315
Fore wing as in male, costa slightly less curved posteriorly, apex more produced,
termen ochreous-tawny. Costal strigulae slender, rather pale, except on costal edge
where they are brown; streaks along veins beyond cell more contrasting, brownish;
paler area beyond cell reduced to a narrow streak between veins 7 and 9; there is
also a pale transverse spot along closing vein; lower edge of cell broadly suffused
with tawny-brown; slight dusting with dark brown indicating first and second
discal spots, terminal fascia narrow, black dot in apex present. Cilia tawny, densely
dusted with purple, a purple supramedian and a slender submedian lines.
Hind wing glossy whitish-golden, veins pale yellowish, extreme apex infuscated.
Cilia concolorous, towards apex clouded with fuscous.
Other females paler, straw-ochreous, one more greyish-fuscous tinged; every
one has the characteristic black apical dot; one specimen (4300) is fasciate: whitish-
ochreous, with strongly reduced strigulae and with a broad dark fuscous-grey
median longitudinal streak.
Male genitalia. Resembling those of the Australian psammitis Turner, but with
differently spined sacculus. Tegumen low and robust. Uncus moderate, top with
corona small. Socius rather large. Vinculum band-like. Valva broad and short,
suboval. Cucullus broad, rounded, sparsely spined. Valvula moderate, with two
lateral spines beyond the corona. Sacculus large, 1/7, concave, with a semicircular
series of 6—7 very strong spines, crossing the disc of the sacculus; punctulate area
moderate, triangularly projecting laterad. Some fine setae scattered over base of
sacculus. Aedeagus rather short, robust.
Female genitalia. Top of abdomen slender and elongate. Eighth segment rigid,
sclerotized, compressed laterally (this does not show in fig. 38, does in fig. 41).
Sterigma simple, a depressed-triangular lobe on each side of ostium, strongly
sclerotized in middle, with a broad transparent upper edge; below each of the
lobes, a rounded ill-defined sclerotization. Ostium not modified. Colliculum, a
moderate, slightly sclerotized tube. Lobus analis slender and long, some wrinkles
below foot of this lobe. Signum small, basket-shaped.
Material studied, in the Transvaal Museum. Males. South Africa, Transvaal:
Mahuba’s Kloof, 1050 m, 14.1.1925 (A. J. T. JANSE), 1 &, holotype, gen. no.
4364. Paratypes, the same locality and date, gen. no. 4361. Marieps Mtn. 3.1.1926
(G. VAN SON), gen. 4406. Pretoria, 4.XI.1909, gen. 4322; 8.XI.1909 (4327);
Waterval-Onder, 10.X.1910 (4319) (A. J. T. JANSE).
In the British Museum. Pretoria, 1.1910 (JANSE), paratype, 4, gen. 3618. In
totali
Females. South Africa, Transvaal: Barberton, 27.XII.1910 (A. J. T. JANSE),
allotype, gen. no. 4306. Paratypes: the same locality and collector, 28.1.1911, gen.
nos. 4300 and 4341. Natal: Sarnia, XI.1913 (Mrs. Burry, Coll. JANSE), gen. no.
4289. Cape Province: Pt. St. Johns, 1-7.1.1931 (A. J. T. JANSE), gen. no. 4480.
(In the Transvaal Museum).
Natal: Weenen, 1.1925 (H. P. THOMASSET, MEyRIcK Collection) 29, gen.
nos. 2526 and 2527. (In the British Museum). In total 9 9.
Fig. 21. Bactra (Chiloides) jansei sp.n., & holotype, genitalia. Fig. 22. The same of 9,
allotype. Fig. 23. The same, bursa
316 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
A rather small species of very variable and therefore confusing appearance.
This material was identified by MEYRICK with stagnicolana ! The females are
slightly larger. The pale hind wings in the two sexes are rather characteristic.
The nearest relative, at present known to me, occurs in New Guinea and
Australia (B. (C.) scalopias Meyr.).
Dedicated to Professor Dr. A. J. T. JANSE, the nestor of the South African
Lepidopterology and collector of most of the present material.
Bactra (Chiloides) adelpha spec. nov.
Figs. 24—26
& 20 mm. Head and thorax fuscous. Palpus rather dark fuscous-grey. Abdomen
pale greyish-ochreous.
Fore wing oblong, dilated, costa curved at base, almost straight posteriorly, apex
moderately pointed, termen sinuate, oblique. Tawny-ochreous, densely dusted and
suffused with fuscous and fulvous, veins streaked with darker, rows of ferruginous
points between veins (superficially the wing seems to be rather evenly fuscous-
fulvous coloured and almost devoid of markings). Costa with a series of very
short dark greyish-fuscous suffused marks; a moderate brownish streak in posterior
part of cell, sharply limited by stigma which is whitish, very slender and fasciate,
actually being a strigula along closing vein, gently convex outwardly; a faint tawny-
ferruginous suffusion along termen, dilated upwards. Cilia purplish.
Hind wing glossy pale grey, on apical 2/3 tinged brownish-fuscous, with a
faint bronze gloss. Cilia concolorous.
Male genitalia. Similar to those of the preceding species (janser), but more
robust, with larger socius and a more defined tuba analis. Valva less broad,
especially cucullus. Sacculus with a fasciate and extended narrow punctulate area,
with at the base a few small spines, and at the top of this area with some five
slender and bristle-like spines; a similar single spine on top of right sacculus, left
sacculus with four more robust apical spines. Aedeagus somewhat longer.
South Africa, Transvaal: Pretoria, 3.XII.1909 (A. J. T. JANSE), 1 4, holotype,
gen. no. 2903.
9 23 mm (allotype). Head and thorax rather deep tawny, slightly suffused
with fuscous and with faint fulvous gloss in certain lights; face and vertex in
middle rather bright ochreous-tawny. Palpus long, pointed, slightly rough along
upper edge, terminal segment almost concealed; dark brown. Abdomen fuscous.
Fore wing elongate, rather narrow, costa curved at base, straight posteriorly,
apex pointed, termen sinuate, oblique. Bronze-brownish, with slight traces of
markings only. Costa suffused with fuscous-purple; an ill-defined broad median
fuscous-purple streak from base to apex; a faint ochreous-yellowish streak from
middle of upper edge of cell to apex; termen slightly mixed with brighter ap-
pearing tawny-orange in certain lights. Cilia (damaged) dark brown, with a series
of whitish round spots along basal half, arranged on opposite ends of veins.
Hind wing light leaden-fuscous, with some yellowish tinge, dorsum as far as
cell sordid pale leaden, without yellowish. Cilia whitish-leaden; along fuscous part,
pale fuscous with a paler base and a fuscous subapical band.
Female genitalia. Sterigma simple, showing only a weak rhomboidal or sub-
Fig. 24. Bactra (C.) adelpha sp.n., 3, holotype, genitalia. Fig. 25. The same of ©, allotype.
Fig. 26. The same, bursa.
pentagonal ostium. This is, however, flanked by characteristic structures, being
semioval impressions, open laterally, with a sclerotized edge that is margined by a
hyaline substance. Colliculum, a slender, simple tube, slightly spindle-shaped, with
several longitudinal folds. Postapophyses not perceptible. Signum moderate, basket-
shaped, rather shallow.
South Africa, Transvaal: Tweefontein (near Bronkhorstspruit), 13.1.1907 (in
JANSE's hand, 1 @,allotype, gen. no. 4476.
The species is much larger and not at all similar to jansei superficially, but
judging from the genitalia, closely allied.
318 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
Bactra (Chiloides) simpliciana Chrétien, 1915
Bactra simpliciana Chrétien, 1915, Ann. Soc. ent. France, vol. 84, p. 302 (larva and pupa,
on Cyperus conglomeratus Rottb.). — Amsel, 1951, Bull. Soc. Sci. Nat. Maroc, vol. 31, p. 68.
Bactra (Chiloides) telaviviana: Amsel, 1951, Beitr. nat. Forsch. S.W. Deutschl., vol. 17, p.
76, fig. 18 (4 N.E. Arabia).
Bactra telaviviana Amsel, 1935, Zoogeographica, vol. 2, p. 46, 123 (nom. nud.). — 1935,
Mitt. Zool. Mus. Berlin, vol. 20, p. 291, pl. 11, fig. 89 (Tel Aviv, Palestine). — 1935,
Veröff. Deutsch. Kol. Uebers. Mus., vol. 1, p. 261, no. 230.
Bactra lactosana Turati, 1922, Atti Soc. ital. Sci. nat. Milano, vol. 61, p. 172 (Cyrenaica;
sex?). Syn. nov.
Bactra pauperrima Turati, 1934, loc. cit., vol. 73, p. 196, t. 3 fig. 24 (Cyrenaica, sex ?).
Syn. nov.
The two species described by COUNT TURATI remained enigmatic, since the
types are either lost or not accessible. All my informations concerning the fate of
TurATI Collection did not provide any results. The collection seems to be in the
possession of the TURATI family, but my inquieries addressed there were never
answered.
The description of the two species and the figure of pauperrima concur in detail
with those of s/mplictana. Therefore it is apparent that we may sink both former
names as synonyms of the latter, the present species being quite characteristic.
Bactra (Chiloides) sordidata spec. nov.
Fig. 27
& 18 mm. Head and thorax grey-fuscous. Palpus moderate, strongly dilated,
triangular; dark fuscous with a pale greyish median streak, terminal segment very
short, almost concealed, blackish. Abdomen fuscous.
Fore wing oblong, rather broad, costa gently curved throughout, apex obtusely
pointed, termen gently sinuate, little oblique. Glossy purplish-fuscous, rather dark,
slightly dusted with pale grey. Markings dull and therefore contrasting, ferru-
ginous-tawny, more or less olive-tinged, limited (or partly rubbed?). Costal
strigulae numerous, narrow and oblique, anteriorly not reaching cell, posteriorly
some continued so as to form three or four more or less complete curved lines,
running across wing between cell and termen, ending in tornus and lower part of
termen; anterior part of wing except costa less densely strigulated, interrupted into
short fragments; first spot small, dark fuscous, oval, in’ fold well beyond 1/3;
a roundish suffused ferruginous spot filling out space just beyond cell, another
similar but smaller end transverse mark halfway between preceding and apex.
Cilia (imperfect) ferruginous-fuscous.
Hind wing pale grey with a pinkish-bronze gloss, dorsal third whitish. Cilia
concolorous but paler.
Male genitalia. Very similar to those of rhabdonoma, differing as follows.
Socius larger. Vinculum broad and flattened. Cucullus much narrower, not at-
tenuated, top broadly rounded. Valvula with a smaller corona, stalk distinctly
extended in middle, with only a couple of spines. Sacculus broad and concave, but
A. DIAKONOFF : African species of Bactra 319
simple, top truncate, without spines. Punctulate area slightly elongate. Juxta larger.
Caulis longer. Aedeagus thicker, less curved and sclerotized, slightly shorter;
cornuti, two adjacent dense patches of long spines.
South Africa, Natal: St. Lucia Lake, 18.X11.1932 (H. W. BELL MARLEY), 1 4,
holotype, gen. no. 4403. Unique.
Bactra (Chiloides) tylophora spec. nov.
Figs. 32—34
& 21 mm. Head whitish-ochreous, sides of vertex infuscated. Palpus rather
long, slender at base, abruptly triangularly expanded posteriorly, edge rough;
terminal segment short, obtuse, semiconcealed; ochreous-whitish, a dark grey
oblique suffused median band, on lower angle of top of apical tuft this band
becoming black; terminal segment blackish, tip pale. Thorax fuscous (greasy),
tegula pale ochreous dusted with dark fuscous. Abdomen light fuscous.
Fore wing suboblong, broadest at 3/4, costa gently curved throughout, apex
pointed, termen slightly sinuate, oblique, long. Pale ochreous, partially touched
with pale yellow, sharply strigulated and marked with dark fuscous. Base of costal
edge slightly infuscated; anterior half of costa with numerous fine strigulae
reaching to cell, more or less interrupted; posterior half with much more oblique,
almost horizontal strigulae and with indistinct pairs of pale marks, obscured by
numerous very short dark strigulae; an irregular streak of coarse dark fuscous
marbling below cell from about 1/, to lower angle; dorsum dark-marbled, its
edge dark dotted; a dark fuscous suffusion along termen, reaching beyond middle
of distance towards cell, above reaching to apex, below to lower part of termen;
less obscure ground colour forming a streak from middle of cell to costa just
before apex; this streak containing sharp and fine lines along edges of cell and
along most veins; an area between posterior half of costa and dorsum similarly
unobscured, pale ochreous; stigma snow-white, cutting dark line along lower par-
ting vein and vein 6 in two parts; an interrupted line along lower half of termen.
Cilia pale ochreous, an antemedian, a postmedian line and tips of cilia fuscous.
Hind wing rather dark fuscous-grey, paler towards base, with a faint bronze
gloss. Cilia pale fuscous, mixed with whitish, a darker fuscous subbasal band.
Male genitalia. Tegumen moderate. Uncus rather short and broad. Socius large.
Valva moderate, cucullus broad, simple, suboval, costal edge hardly sinuate; valvula
rather short, stalk slender, top broad; sacculus deeply concave, devoid of spines,
towards smoothly rouded top a submarginal longitudinal patsch of minute hairs;
only median posterior angle of the inner edge of sacculus extended into a strong
characteristic process, with top blunt and slightly laminate; an oblique transverse
band of small hairs across sacculus above its base. Juxta robust, rather small. Vincu-
lum normal. Aedeagus broad, rather short, bent. Cornuti, several (11) strong
spikes and a patch of smaller spines.
South Africa, Transvaal: Waterval-Onder, II.1912 (BONNEKAMP, Coll. A. J. T.
JANSE), 1 4, holotype, gen. no. 4292. Orange Free State: Bloemfontein, 23.XI.-
1925 (H. E. IRVING), 1 &, paratype, gen. no. 4365.2 ¢.
A robust species with finely dark-marbled and lined fore wings. Allied with
320 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
A. DIAKONOFF : African species of Bactra 321
fasciata where a process of the internal edge of sacculus is already developed but
short.
9 20 mm. Head, palpus and thorax pale tawny, but brighter than in male,
palpus with an oblique dark grey streak laterally. Fore wing as in male but costa
a trifle less curved. Whitish-ochreous, strigulated with pale ochreous-tawny, on
anterior half of wing markings interrupted in series of numerous short strigulae,
on posterior half markings obliterate, being obscured by a brighter tawny suffusion
from cell to termen, between veins 4—7; costal markings minute, dark brown,
short, first discal spot indicated by a dark fuscous vertical strigula, second discal
spot, by a suffused fuscous rounded spot below lower angle of cell; a whitish short
streak along bases of veins 3—5, veins themselves finely brown; a terminal streak
and a suffusion below apex, brownish.
Hind wing slightly paler, especially towards apex where it is tinged ochreous.
Female genitalia. Sterigma, a moderate but strong rounded sclerite, with the
upper edge truncate, the lower narrowed and truncate or folded over. Ostium in
centre of this sclerite, strongly sclerotized, rather small, colliculum, a narrow rather
long tube, appearing as a narrow sclerotized band on each side, each band with
hyaline edges. Signum basket-shaped.
South Africa, Transvaal: Waterval-Onder, 23.XI.1910 (A. J. T. JANSE), 1 9,
allotype, gen. no. 4348. 1 specimen.
The female is more ochreous tinged than the male and less sharply marked, but
otherwise very similar.
Bactra (Chiloides) endea spec. nov.
Figs. 28—31
:9 11—14 mm (holotype 12 mm). Head and thorax creamy, vertex pale tawny.
Palpus strongly dilated, with a rough edge; creamy, infuscated except at apex,
terminal segment exposed, slender, fuscous. Abdomen glossy whitish.
Fore wing suboval, costa rather curved throughout, apex pointed, termen gently
sinuate, considerably oblique. Creamy-white, minutely strewn with brown (tips of
scales!), markings narrow, well-defined, dark brown. Anterior part of costa with
a series of short marks, posterior part with about five very distinct oblique and
long streaks, on costa dark brown, below costa becoming olive-tawny and
narrowed; these streaks alternating with minute dark brown transverse marks;
basal patch indicated by a curved, slightly outwards-oblique line, from before 1/4
of costa to well beyond 1/3 of dorsum, oblique above, becoming less oblique
below, in fold including the dark brown conspicuous first discal spot; this spot
elongate, acute anteriorly, tolerably rounded posteriorly; second discal spot large,
tawny touched with olive, crescentic-suboval, arranged longitudinally, edges some-
what suffused; anterior arm of this spot connected with the first costal streak
originating from slightly before middle of costa, posterior arm very short; stigma
creamy, elongate-oval, not conspicuous; apical streak brownish mixed with purple
Fig. 27. Bactra (C.) sordidata sp.n., & holotype, genitalia. Fig. 28. B. (C.) endea sp.n.,
9 holotype, sterigma and ovipositor. Fig. 29. The same, bursa. Fig. 30. The same, ® para-
type, gen. no. 3520, sterigma and ovipositor. Fig. 31. The same, part of bursa with signum.
322 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
dusting, extended into an elongate shade, becoming darker posteriorly; an
elongate black dot in apex; terminal edge with a tawny marginal streak; dorsum
with a series of slender dark brown transverse marks. Cilia creamy, densely dusted
with purplish, so as to form several narrow darker lines.
Hind wing creamy with a golden gloss, posterior half gently infuscated. Cilia
concolorous, along termen and around apex infuscated, and with a subapical
fuscous band. The paratypes are all rather uniform and similarly marked; only one
specimen (3651) is moderately infuscated all over the wing.
Female genitalia. Sterigma very delicate, only lightly sclerotized. Ostium bursae
and colliculum trumpet-like, top dilated laterally and rather mushroom-shaped,
under this “hood”, a series of semicircular folds. Edge of seventh segment mod-
ified into two oval plates (less distinct in holotype). Ductus bursae abruptly nar-
rower than colliculum. A small sclerite is present representing a cestum. Signum,
rather small, basket-shaped.
West Africa, Gambia: Bathurst, XI.1884 (CARTER), 1 9, holotype, gen. no.
3652; the same, 1884, 1 9 paratype, gen. no. 3656; the same XI.1884, 1 9,
paratype, gen. no. 3651; the same, XI-XII.1885, 2 9, paratypes, gen. no. 3617,
3636. Nyassaland: Mt. Mlanje, 28.1.1914 (S. A. NEAVE), "Bactra pythonia Meyr.,
teste Meyr. 683”, 1 9 , paratype, gen. no. 3520. 6 9.
The species is extremely close to B. (C.) angulata Diak., from Borneo, Java,
Moluccan and Palau Islands. The slender species have the same facies and mark-
ings. The genitalia, however, differ sufficiently to justify the separation of endea.
The ostium bursae in angulata is stronger, at top more compressed so as to form
a median and two lateral spout-like folds, plates at the foot of ostium are standing
upright and are much more sclerotized, the caudal edge of the eighth segment
is more distinctly modified.
The males are so far unknown.
Bactra (Chiloides) rhabdonoma spec. nov.
Figs. 35—37
& 13—17 mm (holotype 17 mm). Head creamy with a faintest pinkish tinge
on vertex. Palpus moderate, strongly dilated, fuscous-grey. Thorax creamy, tegulae
touched with pinkish, with a pinkish-lilac gloss in certain lights. Abdomen fuscous.
Fore wing pointed-suboval, rather broad, costa considerably curved throughout,
apex pointed, termen hardly sinuate, oblique and long. Pale creamy-ochreous,
anterior half faintly pinkish, posterior yellowish tinged. Costa with numerous
oblique dark strigulae, anteriorly dark brown and short, along more than posterior
half becoming extended by slender tawny appendages; interspaces anteriorly grey,
posteriorly forming the usual paired silvery-white marks; a subcostal streak of pale
ochreous ground colour, not obscured or marked, extending from base of wing
to apex, extending downwards anteriorly to middle of cell, posteriorly sharply
delimited by apical streak; first discal spot large, very irregular, fuscous, connected
by some irroration with the second discal spot; this spot conspicuous, deep brown,
semicircular, top of anterior arm extended basad by a short strigula; stigma some-
what irregular, oblique, ochreous-white; apical streak dark brown, gradually dilated
A. DIAKONOFF : African species of Bactra 323
Fig. 32. Bactra (C.) tylophora sp.n., 6, holotype, genitalia. Fig. 33. The same, 9, allotype,
sterigma and ovipositor. Fig. 34. The same, bursa
324 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
posterad, lower edge ill-defined, merging into grey suffusion occupying entire
termen as far as cell; this suffusion mixed with tawny and transversely strigulated
with brownish, but very faintly; a complete band and a subbasal, a postmedian,
and an apical black lines.
Hind wing whitish-ochreous or whitish-tawny, pale, slightly infuscated towards
apex. Cilia whitish, infuscated except towards dorsum, with a whitish basal line.
Male genitalia. Tegumen and uncus normal, moderate. Socius rather small.
Vinculum erected, V-shaped. Valva with a broad cucullus, costa rounded and pro-
minent beyond base, top narrowed and rounded, a rounded moderate prominence
in the middle of lower edge. Valvula rather short but broad, corona, stalk with a
slightly oblique dense row of spines. Sacculus thickened and concave, rather simple,
outer edge with fine bristles, top rounded-truncate, with a small patch of rather
short spines, punctulate area oval, moderate. Juxta rather small. Caulis developed.
Aedeagus sclerotized, robust and long. Cornuti, some 6 scattered moderate spines,
and a subapical very large patch of extremely dense spines.
South Africa, Transvaal: Nelspruit (C. J. S.), XII.1917 (in MEyRICK’s hand),
“Bactra stagnicolana Zell.”, 1 &, paratype, gen. no. 3469 (in the British Museum).
The same locality, I1.1918 (H. G. BREIJER), 2 &, paratypes, gen. no. 4242 and
4244. Marieps Mtn., 10.XII.1925 (G. VAN SON), 1 4, holotype, gen. no. 4286.
The same locality and collector, 5-8.XII.1925, 9 4, paratypes, gen. nos. 4388,
2590450115752
2 11.5—22 mm (allotype 15.5 mm). Head and thorax whitish-ochreous, thorax
faintly spotted with grey, a grey subapical spot on each tegula. Palpus abruptly and
strongly dilated, roughish; pale ochreous, median segment with a black point in
middle of inner edge and a fuscous shadow opposite this on the outer edge. Abdo-
men light fuscous.
Fore wing suboval, rather broad, costa gently curved throughout, apex pointed,
termen straight, oblique. Pale grey or whitish with a distinct bluish-leaden gloss.
A longitudinal supramedian streak from base to apex, dull creamy-white. Other
markings dark fuscous and pale tawny. Costal strigulae distinct, moderate, rather
distant from each other (about 9 in total), alternating with minute dark
marks, both dark brown; median strigula faintly reaching to second discal spot,
penultimate strigula reaching to termen; a black dot in apex; first discal spot
distinct, fasciate, transverse, direct, beyond 1/3, blackish, slightly mixed with
olive-tawny; this spot preceded by a series of three similar, blackish spots; second
discal spot incomplete, only inner arm being present and forming an oblique
blotch, dark fuscous mixed with olive-tawny; stigma suffused, white, small; apical
streak incomplete, formed by a small, ill-defined tawny spot beyond cell and a
larger one of irregular fuscous suffusion, before termen; ocellar area centred with
an elongate transverse dot, cut in two by vein 3; a dark fuscous terminal fascia,
attenuated downwards and interrupted between veins 5—7. Cilia dark fuscous,
dusted with white.
Hind wing pale grey, extreme apex suffused with darker grey. Cilia concolorous.
Female genitalia. Seventh sternite sclerotized, forming two large oval plates with
a hyaline outer edge; their caudal edge is wrinkled, two areas of fine wrinkles
flanking the ostium. Ostium small, triangular, simple. Colliculum, a moderate,
A. DIAKONOFF: African species of Bactra 325
Fig. 35. Bactra (C.) rhabdonoma sp.n., &, holotype, genitalia. Fig. 36. The same, 9 allo-
type, sterigma and ovipositor. Fig. 37. The same, bursa
326 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
slightly narrowed tube. Cestum, a small oval sclerite. Signum, a shallow basket.
A variable series, as to the colouring as well as to the size. Larger specimens
are more tawny-tinged, smaller, pale greyish. The markings are rather similar and
appear superficially as a narrow median streak interrupted by the stigma, a very
narrow suffusion along costal margin, and the minute ocellar dot.
Material studied. In the British Museum. South Africa, Transvaal: Waterval-
Onder, 18.XI.1910 (A. J. T. JANSE) (label in MEYRICK's hand, in MEYRICK
Collection), 1 9, gen. no. 3613. In the Transvaal Museum. Transvaal: Barberton,
17.1.1911 (A. J. T. JANSE), 1 ©, allotype, gen. no. 4316. The following are all
Q paratypes. Natal: Umkomaas, 10.1.1914, gen. no. 4374; 17.1.1914, gen. nos.
4287 and 4457; 24.1.1914, gen. no. 4456 (all by A. J. T. JANSE). Transvaal:
Marieps Mtn., 8.XII.1925 (G. VAN SON), gen. no. 4398.6 9.
The species apparently is intermediate between the coronata and the venosana
groups, linking them together; the male seems closer to the latter species, while the
female seems related to coronata 9.
Bactra (Chiloides) stagnicolana Zeller, 1852
Figs. 38—41
Bactra stagnicolana Zeller, 1852, Lepid. Micropt. Caffr., p. 84, (’’Caffraria”). — Meyrick,
1913, Ann. Transv. Mus., vol. 3, p. 275 (siccella Walk., synonym). — 1920, Voyage Allaud
& Jeannel, Lep., p. 63 (E. and S. Africa). — Diakonoff, 1958, Ent. Tidskr., vol. 78, suppl,
p. 77, text fig. 7, tab. fig. 7 (type redescr., gen. & fig.).
Tortrix stagnicolana: Walker, 1863, List Lep. Het. Brit. Mus., vol. 28, p. 326 (S. Africa).
?Ancylolomia siccella Walker, 1866, l.c., vol. 35, p. 1750 (2, Cape).
?Aphelia lanceolana Walsingham, 1881 (nec Hübner), Trans. Ent. Soc. Lond., p. 231
(Natal, Spring Vale, Dec., March, April)
Distribution. South and East Africa.
It is not possible to ascertain whether "siccella’ is a synonym of stagnicolana,
because the unique type specimen of WALKER seems to be lost. It could not be
located neither in the British Museum nor in the Oxford Museum. Neither am I
able at present to make out the identity of the material which LORD WALSINGHAM
studied in 1881. At that time he regarded Bactra all over the world as belonging
to a single species, lanceolana; the three specimens he saw from Spring Vale may
have been as many different species.
For a redescription and figure of ZELLER’s male holotype may be referred to my
earlier paper (1958).
The species is variable; the following types may be discriminated.
(1). Maculate type. The ground colour pale purplish-grey, the markings fer-
ruginous-fuscous. The costal strigulae are numerous and rather slender. The first
discal spot rounded, fuscous, across fold at 1/3; the second distal spot broad,
semioval, along lower angle of cell; pretornal mark well defined, moderate,
transversely-oval, closer to preceding than to tornus; apical streak almost black,
well-defined, running from middle of wing across stigma and more or less
obscuring it, to apex; sometimes there is a distinct circular thick brown line from
middle of vein 8 to and along lower third of termen; sometimes veins beyond cell
more or less narrowly dark-streaked. This is the common type, in males as well as
in females.
A. DIAKONOFF : African species of Bactra 327
1 i
Fig. 38. Bactra (C.) stagnicolana Zell, & holotype, genitalia. Fig. 39. The same, ® allotype,
ovipositor and sterigma. Fig. 40. The same, bursa
328 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
(2). Suffused type, is similar to preceding, but with the wing evenly suffused
with dark grey, so that all discal markings disappear and sometimes only marginal
strigulae remain; stigma minute, but distinct. Also common in males, less common
in females. The holotype belongs here.
(3). Unicolorous type, only females. Pale ochreous, sometimes more or less
darker infuscated along edge of wing, with traces of costal strigulae; sometimes
also a faint infuscation of centre of disc.
(4). Vittate type, only females. Ground colour pale ochreous, costal markings
very short and faint, apical streak dark brown, including the clear white stigma,
and connected with the base of wing by a fuscous suffusion along middle of disc;
pretornal mark reduced to two short horizontal black strigulae.
Hind wing invariably whitish-ochreous, pale, slightly infuscated towards apex,
seldom gently suffused with pale grey all over. Hind wing in female paler than
in male.
The female has never been described. Therefore a female neallotype is selected
herewith, gen. slide no. 3798. This specimen is of the vittate type, 20 mm, and is
labelled thus: Pretoria, 21.XII.1911 (A. J. T. JANSE).
Male genitalia. Tegumen rather broad, lower part rounded, top high and slender.
Uncus moderate, slender. Socius normal, rather small. Tuba analis large and
distinct. Vinculum strongly dilated at the sides, normal below. Valva broad;
cucullus characteristic, broad, with an obliquely rounded top; costa broadly prom-
inent in middle; lower edge of cucullus above the primary incision with a dense
corona of long spines and bristles; valvula with a broad top and a moderate stalk;
sacculus deeply folded and concave, top forming a short, prominent and serrulate
crest; a strongly sclerotized subtriangular tumescence subapical on the inner edge
of sacculus, crowned with a group of 5—6 moderate spines, anterad from these a
patch of small hairs. Aedeagus peculiar and therefore very characteristic, sclerotized,
base spherical, above base aedeagus laterally flattened, broad, narrowed towards
top, upper edge with a postmedian prominence bearing three small teeth; ventral
side of aedeagus formed by a separated slender sclerite, at top bent sideways and
denticulate.
Female genitalia. Lobus analis long and rather narrow, postapophyses long.
Eighth segment erected-conical, moderately sclerotized; anapophyses moderately
long. Lamella postvaginalis shaped as a moderate transverse band, being the caudal
portion of the eighth sternite. Colliculum attached to its middle, very large,
spindle-shaped, with slightly strengthened upper wall and below middle thickened
walls, in the middle a narrow, strongly sclerotized, lip-like and sinuate transverse
fold. Ostium membraneous, rather wide. Ductus bursae wide, moderately long.
Corpus bursae large, with a finely scobinate wall. Signum moderate, basket-shaped.
(Slide figured, 3798).
Material studied. In the British Museum. Comoro Islands: Mayotte (L.),
V.1911, 2 & (3523, 3525), 2 9 (3522, 3524). Nyassaland: Fort Johnston, (R.)
III.1906, 1 :8 (3616). South Africa, Cape Province: Ceres (R. E. TURNER),
III.1925, 1 :9 (3641). Transvaal: Barberton, VI.1911 (A. J. T. JANSE), 1 2
(3463). Natal: Weenen, XII.1926 (H. P. THOMASSET), 1 $ (3626). Madagascar:
Ampoza, IX.1929 (E. J. WHITE) (B.M. 1930—26), 19, gen. no. 3075.
A. DIAKONOFF : African species of Bactra 329
Fig. 41. Bactra (C.) stagnicolana Zell, & holotype, part of genitalia, showing the compres-
sed aedeagus (after Diakonoff, 1958). Fig. 42. B. (C.) aletha sp.n., &, holotype, genitalia
In the Transvaal Museum. Southern Rhodesia: Bulawayo, Zimbabwe. Transvaal:
Pretoria. Natal: Nkwaleni, Magude, New Hanover, Karkloof, Umkomaas. Cape
Province: Willowmore, Kokstad, Grahamstown, Stella Bush. From October untill
June (A. J. T. JANSE, C. J. SWIERSTRA, F. SIMONS, HARDENBERG, C. G. C.
DICKSON, G. C. CLARK, J. OMES-COOPER, VAN DER MERWE). 18 ¢, 20 9.
Gen. nos. & 3790, 3798, 3804, 4232—33, 4237, 4253, 4267, 4294, 4337, 4349,
4351, 4358, 4405—6, 4644 T.M. 2, 3801, 3805, 4235, 4249, 4250, 4251, 4277,
4297, 4343, 4450, 4471, 447374, 4478, 4481.
In Leiden Museum. Mauritius, Moka, 1 ¢, gen. no. 2880; Mauritius, 1 ¢,
gen. no. 2892 (J. VINSON).
330 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
A big species, often conspicuously dark-coloured (melanistic). Judging from the
male genitalia, related with venosana.
Bactra (Chiloides) punctistrigana Mabille, 1900
igs GOE)
Bactra punctistrigana Mabille, 1900, Ann. Soc. ent. France, vol. 68 (1899), p. 751 (Ma-
dagascar). — Viette, 1947, Mem. Inst. Sci. Madagascar, ser. A, vol. 1, p. 54. — 1954, ibid.,
ser. E, vol. 5, p. 379 (lectotype select.). — Diakonoff, 1959, Rev. franc. Ent., vol. 26, p.
177, figs. 20—21, t. 2 fig. 10 (type redescr., fig.).
Distribution. Madagascar.
For the description of the female type specimen may be referred to my above
cited paper (1959). That specimen is melanistic, while the female specimen at
hand (gen. no. 4234) is brighter coloured and marked. It may be redescribed thus.
? 20 mm (right pair of wings missing). Head pale ochreous, thorax greyish-
tawny (palpi missing). Abdomen light fuscous. Fore wing slightly broader than
in the type, dilated, costa gently curved, apex pointed, termen distinctly sinuate,
little oblique. Pale ochreous, rather densely suffused with fuscous-grey, except a
pale streak between cell and costa; costa narrowly dark fuscous, transverse markings
pale ochreous; median longitudinal fascia interrupted anteriorly, grey-fuscous and
cloudy along cell, ill-defined, posteriorly forming a dark brown, well-defined
apical streak; stigma, small white, rounded; a dark brown terminal fascia, rather
broad, attenuated below, almost interrupted between veins 5—6; a longitudinal
dark brown mark in centre of ocellus. Cilia pale tawny with a whitish basal line
and a postmedian dark line, on lower part of termen and in tornus cilia uni-
colorous light tawny.
Hind wing ochreous-whitish with a silky gloss. Cilia glossy, creamy.
Female genitalia are described in my 1959 paper. They show such a close
resemblance to those of venosana that at first I regarded MABILLE’s species as a
subspecies of the former, no males being available at the time. Now I am satisfied
that punctistrigana is a perfectly distinct species, although closely related with
venosana, as the genitalia of the two sexes show.
Material studied. South Africa, Southern Rhodesia: Zimbabwe, 6-10.1V.1934
(A. T. J. JANSE), 1 9, gen. no. 4234. Madagascar: Sumbiran Plantation, 1911
(R.), Meyrick Collection (in the British Museum), 1 9 gen. no. 3615.
& neallotype, 19 mm. Head and thorax dark fuscous, metathorax silvery-grey.
Palpus rather long, strongly dilated triangularly, densely and tolerably smoothly
scaled; dark fuscous, terminal segment almost entirely concealed. Abdomen glossy
greyish-white.
Fore wing oblong-subtruncate, costa rather strongly curved along anterior half,
straight along posterior, apex subobtuse, termen hardly concave, almost straight,
little oblique. Dark fuscous, markings faint, ferruginous. Costa with numerous
little curved transverse strigulae edged on both sides and accentuated with faint
leaden-purplish; costal strigula at 4/5 of costa longer than other, straight, almost
to termen below apex; a ferruginous terminal streak, along extreme margin of
A. DIAKONOFF : African species of Bactra 331
wing finely dusted with whitish. Cilia with basal half dark fuscous, apical half
paler fuscous, base with a narrow pale line, a postmedian blackish line.
Hind wing glossy pale greyish, posterior half slightly infuscated, apex darker
suffused with grey. Cilia glossy pale grey, base with a narrow dull ochreous line,
followed by a dark grey band.
Male genitalia. Of the venosana type. Tegumen high and rather narrow. Socius
large. Tuba analis perceptible. Vinculum, a strong, broad band, mesially rounded-
prominent, smooth. Valva broad. Cucullus broad, broadest above base, top oblique
and attenuated, costa being strongly sinuate. Sacculus extremely large, extended and
deeply impressed from the outside, so that it is not possible to open and flatten
both valvae without dislocating them; outher surface smooth except for an ex-
tremely dense basal patch of strong slender spines, upper angle of the inner edge
of sacculus with a short, curved blade; inner surface with a narrow strip of
punctulations (in left valva showing through, in right plainly visible), and a
continuation of the basal patch of spines, partly on an elevated short ridge. Juxta
and caulis strong. Aedeagus long and curved, with a slender longitudinal sclerite
dorsally, on the right side beset with strong cornuti.
Material studied. Central Madagascar: Imerina Plateau, Tsimbazaza Park, 1200
m, 11.1.1952 (P. VIETTE), 1 &, neallotype, gen. no. 2611. In the Paris Museum.
Unique.
Obviously this is the sex partner of the species of which the female was des-
cribed by MABILLE. The facies, colouring, locality, and above all the character of
the ¢ Q genitalia point in this direction.
Bactra (Chiloides) venosana (Zeller, 1847)
Phoxopteris venosana Zeller, 1847, Isis, p. 738.
Aphelia venosana: Herrich-Schäffer, 1849, Syst. Bearb., vol. 4, p. 244.
Bactra venosana: Rebel, 1901, in Staudinger & Rebel Catal., vol. 2, p. 113. — Kennel, in
Spuler, 1910, Schmett. Eur., vol. 2, p. 273. — Kennel, 1910, Pal. Tortr., p. 472, t. 18 fig.
73. — Diakonoff, 1956, Zool. Verh. no. 29, p. 33, figs. 31—33.
Bactra (Chiloides) venosana: Diakonoff, 1956, Zool. Verh., no. 29, p. 33, figs. 31—33.
— 1959, Bijdr. Dierk., part 29, p. 184 (Palestine, Sardinia). — 1963, Ann. Mus. Wien, vol.
67 (in press; banosii, truculenta syn.).
Bactra truculenta Meyrick, 1909, Journ. Bombay Nat. Hist. Soc., vol. 19, p. 586. — 1922,
Exot. Microl., vol. 2, p. 521. — Meyrick in Caradja, 1934, Iris, vol. 48, p. 33. — 1935,
Mater. Microl. chin. Prov., p. 57. — Diakonoff, 1950, Bull. Brit. Mus., Ent., vol. 1, p. 289,
t. 5 fig. 16, t. 7 fig. 30. 1956, Zool. Verh., no. 29, p. 27, figs. 28—30. Syn. nov.
Bactra (Chiloides) truculenta: Diakonoff, 1956, Zool. Verh., no. 29, p. 27, figs. 28—30.
— 1959, Bijdr. Dierk., part 29, p. 184.
Bactra scythropa Meyrick, 1911, Proc. Linn. Soc. N.S. Wales, vol. 36, p. 284.
Bactra geraropa Meyrick, 1932, Exot. Microl., vol. 4, p. 147.
Bactra banosii Gozmany, 1960, Ann. hist.-nat. Mus. Hung. vol. 52, p. 416, figs. 3A-D.
Syn. nov.
Distribution. Northern Africa, Southern Europe, Southern Asia, Pacific, Australia.
Bactra (Chiloides) trimera spec. nov.
Figs. 55—57
3 13—16 mm (holotype 16 mm). Head, palpus and thorax creamy-whitish,
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
332
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A. DIAKONOFF : African species of Bactra 333
median segment of palpus with an oblique rather narrow light fuscous transverse
fascia; lower edge of palpus roughish. Abdomen fuscous.
Fore wing elongate, dilated, broadest beyond 3/4, costa gently curved at
extremities, rather straight in middle, apex obtusely pointed, termen gently sinuate
above, little convex below, oblique. Glossy whitish, suffused with pale tawny.
Markings dark fuscous. Costal edge dark fuscous, broken in numerous short marks
by pairs of thick pale transverse strigulae, along posterior portion of costa becoming
silvery; first discal spot longitudinally extended, ill-defined, irregular-oblong or
fasciate, from well beyond base towards second discal spot, but twice interrupted
before latter; second discal spot rather well-defined, V-shaped, anterior arm
broader; stigma moderate, white; apical streak short, black, from below apex
halfway towards cell, edged above with grey; some grey suffusion below this
streak, reaching to cell and more or less extending over terminal part of wing;
ocellar area centred with a wedge-shaped, ill-defined mark parallel to termen, cut
twice by black; apex with a dark brown dot; a dark brown terminal line: a
transverse narrow preapical line, from penultimate dark costal space to termen
below apex. Cilia dark fuscous dusted with pale ochreous, with three dark lines.
Hind wing creamy-white, more or less suffused with pale grey, more so on
posterior half, deeper suffused towards apex.
Male genitalia. Tegumen high. Uncus moderate, curved. Socius rather large.
Vinculum moderate, smooth. Cucullus long, moderately broad, bent and slightly
sinuate over its whole breadth at 1/3 of length, not narrowed, top gradually
rounded; valvula rather long, stalk swollen in middle, top bent, corona rather
small; sacculus concave, strongly flattened dorso-ventrally, top subobtuse with
strong inequal spines, three right, two left, an external submarginal row of short
and slender bristles, internal flattened side with a large patch of long spines at
and above outer lower angle and a second smaller but much denser patch of
smaller spines proximad of the first; haired area from below apex of sacculus to
spine patches, narrow above, in a shallow depression, flanked by two rounded
prominences of the inner edge of sacculus; this hairy area much extended below.
Aedeagus moderate, strongly sclerotized, moderately attenuated and curved,
without cornuti.
The rather pale species makes a superficial impression of having three longitu-
dinal slender dark stripes: costal, median, and dorsal. The five males available are
rather uniform, one somewhat more distinctly marked than the other. Male no.
4264, however, has a fine and sharp dark fuscous marbling upon pale ground,
with veins marked by fine dark lines (”tigroid’’ type of markings).
Material studied. South Africa, Transvaal: Minastone (= Louis Trichardt), 16-
25.11.1919 (H. G. BREIJER), 1 &, holotype, gen. no. 4240. Paratypes: Transvaal:
Waterval-Onder, 10.XI.1910, 1 4, gen. no. 4321. Natal: Nkwaleni, 10.1.1916,
1 4, no. 4463 (both: A. J. T. JANSE). Transvaal: Marieps Mtn., 3.1.1926 (G.
VAN SON), 1 &, no. 4389; Cape Province: Umtata, 20 miles E, 8-11.1.1931 (A.
J. T. JANSE), 1 4, no. 4264. 5 8.
Fig. 43. Bactra (C.) punctistrigana Mab., 4 neallotype, genitalia. Fig. 44. The same, @
holotype, sterigma and ovipositor. Fig. 45. The same, bursa. (Figs. 44 and 45 after
Diakonoff, 1959).
334 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
® 18—20 mm (allotype 18 mm). Head, palpus and thorax pale ochreous-
tawny, palpus spotted with pale grey laterally. Abdomen fuscous.
Fore wing oblong, moderately broad, little dilated, costa gently curved troughout,
apex subobtuse, termen tolerably straight, oblique. Pale ochreous-tawny, in disc
moderately suffused with that colour; costal strigulae numerous, on costal
edge dark brown; those along posterior part of costa tending to form sinuate
transverse strigulae, anterior reaching to dorsum before tornus, posterior strigulae
running to termen; a small pale spot on costa before apex; first discal spot before
1/3, arrowhead-shaped, preceded by several irregular smaller spots along and
around fold; second discal spot very narrow, fasciate and transverse, with a strigula
along closing vein, stigma absent; a suffused tawny terminal streak attenuated
above, edged along extreme margin of wing with dark brown dusting; a series
of dark brown dots along dorsum. Cilia dark fuscous, finely dusted with pale
ochreous, with a narrow whitish basal line.
Hind wing glossy whitish-golden, cilia concolorous.
Female genitalia resembling those of the Asiatic optanias. Sterigma strongly
sclerotized, forming a convex subtriangular, rather three-lobed plate; also the eighth
tergite is sclerotized. Ostium bursae rather ill-defined, its outer rim sclerotized and
curved. Colliculum asymmetrical, strong, clavate. Signum rather small, basket-
shaped.
Material studied. South Africa, Natal: Mtunzini, 30.X1.1953 (L. Vari), allo-
type, gen. no. 4378. Umkomaas, 15.1.1914 (A. J. T. JANSE), 1 ®, paratype, gen.
4305. Cape Province: Kleinmond, IV.1942 (V. G. Dickson), 1 9, paratype, gen.
4296. Pt. St. Johns, 1-7.1.1931 (A. J. T. JANSE), 1 2, paratype, gen. 4375.
The female is rather larger than the male. The genital characters strongly sug-
gest that the sexes belong together, in analogy with the nearest allied Asiatic
species, Bactra (C.) optanias Meyr.
Bactra (Chiloides) aletha spec. nov.
Fig. 42
& 11.5 mm. Head, thorax and palpus fuscous (rubbed!). Abdomen sordid
whitish.
Fore wing oblong, rather narrow, costa gently curved anteriorly, hardly curved
posteriorly, apex moderately pointed, slightly projecting, termen distinctly sinuate,
little oblique. Pale fuscous, clouded with darker (strongly rubbed). Cilia lacking.
Hind wing pale greyish. Cilia missing.
Male genitalia. Belong to the type of boschmai Diakonoff. Tegumen high.
Uncus short and rather broad. Socius moderate. Vinculum strong, V-shaped. Valva
rather slender. Cucullus slender and long, not dilated, costa not prominent, top
rounded; spines small. Valvula rather short, ventral edge with 3—6 rather strong
spines. Sacculus strongly concave, strongly bent rectangularly at base, inner edge
emarginate in middle, strong and partly beset with long bristles, its upper angle
Fig. 46. Bactra (C.) fasciata sp.n., & holotype, genitalia. Fig. 47. The same, paratype,
aedeagus in lateral aspect, gen. no. 3459. Fig. 48. The same, ® allotype, sterigma and
ovipositor. Fig. 49. The same, bursa
A. DIAKONOFF : African species of Bactra 335
336 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
forming an acute strong cusp (not so long as in clarescens or in boschmaz); top
of sacculus with 5—6 strong spines of moderate length; base of sacculus with a
group of robust shorter spines. Juxta and caulis robust. Aedeagus long, curved.
Cornuti apparently absent.
Material studied. West Africa: Kumasi, IX.1911—IV.1912 (J. D. G. SANDERS),
1912—481, 1 3, holotype, gen. no. 3635. Unique. (In the British Museum).
Although the small insect is badly rubbed, the genitalia are distinct and allow
description.
Belongs to the boschmai group of species.
Bactra (Chiloides) salpictris spec. nov.
Figs. 53—54
9 13—15 mm. Head, thorax creamy. Palpus whitish, with a finest dark dusting.
Abdomen pale ochreous.
Fore wing oblong, suboval, pointed, termen hardly sinuate, oblique. Creamy,
with a golden gloss, suffused in disc rather bright ochreous. Markings ochreous
and tawny, becoming dark brown on edges of wing. Costa with the usual strigulae,
very short along anterior half, zigzagged along posterior half, forming some
reticulation or marbling in terminal portion of wing; these markings more or less
interrupted by a longitudinal streak of pale ground colour running along upper
edge of cell and thence to costa before apex; a little distinct median longitudinal
streak from base to termen, formed by ochreous suffusion and markings beyond
cell; terminal area also suffused with pale leaden-grey; first discal spot narrow,
fasciate, transverse, second discal spot small, irregularly semicircular, suffused,
enclosing a very small white stigma; a dark terminal line, an ochreous strigula
before apex; a black dot in apex. Cilia whitish sprinkled with dark brown or
purplish, and with a submedian and an apical dark lines.
Hind wing white, slightly suffused with pale ochreous and greyish towards
apex and costa posteriorly. Cilia whitish, grey opposite apex.
Female genitalia. Eighth sternite, a broad, sclerotized band. Ostium bursae, a
pear-shaped large opening extending almost over the breadth of the band; further
not modified. Colliculum very large and tubular, sclerotized, lower half with a
longitudinal median split, with less sclerotized sides. Signum, a large, shallow
denticulate cup. A pair of peculiar small sclerites apparently belonging to the
anal opening, are visible between the lobi anales.
Material studied. South Africa, Cape Province: Pt. St. Johns, 1-7.1.1931 (A. J.
T. JANSE), 1 2, holotype, gen. no. 4303. Umtata, 20 miles E, 8-11.1.1931 (A. J.
T. JANSE), 1 9, paratype, gen. no. 4366. 2 9.
The paratype is dark, brown-fuscous, a stripe of tawny-ochreous colour from
base to apex, occupying the costal third of wing except the extreme costal edge.
The exact position of the species is uncertain without the knowledge of the
males. It may belong to the boschmai group and then be nearest to aletha.
A. DIAKONOFF : African species of Bactra
337
Fig. 50. Bactra (C. sinassula sp.n., 2, holotype, genitalia
338 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
Bactra (Chiloides) fasciata spec. nov.
Figs. 46—49
& 15.5 mm. Head pale ochreous, face and vertex roughly scaled, scales divided
along middle of face. Palpus thickened, obtuse; pale ochreous above and internally,
dark fuscous elsewhere. Thorax pale ochreous with two pairs of faint fuscous
marginal spots, tips of tegulae fuscous. Abdomen pale ochreous, anal tuft dark
fuscous-grey.
Fore wing elongate-suboval, rather broad, dilated, broadest at 2/5, costa
considerably curved, strongly at base, apex moderately pointed, termen gently
sinuate, concave above, oblique. Pale leaden-grey mixed with whitish, partially
suffused with pale ochreous. Anterior half of costa darker leaden-grey, with several
slender dark fuscous strigulae edged with ochreous suffusion; posterior half of
costa with four pairs of oblique white strigulae, ultimate pair longest, followed
by a vertical single white bar and then by a white smaller triangular dot just
before apex; these white markings converging in one point below costa; anterior
half of wing suffused with pale tawny-ochreous and mixed with some deeper
tawny spots; first discal spot blackish-brown, bent, transverse, across fold just
beyond 1/3, obtusely pointed posteriorly and rather irregular; second discal mark
fasciate, from middle of wing, encircling lower half of stigma, thence to apex,
thus giving rise to a conspicuous apical streak; stigma transverse, conspicuous;
apical streak edged above by a strongly contrasting white streak, from 2/ of disc,
gradually dilated to preapical costal white spot, faintly traversed by minute fer-
ruginous interneural lines and by two transverse similar lines from costa between
the second and third and third and fourth pairs of white strigulae; the latter two
lines gradually curving across wing to 2/3 and 1/, of termen, respectively, merging
there in a dark brown terminal line; ocellar area large, oval, leaden-grey centred
with a brown obliquely-oval dot. Cilia dark grey, slightly dusted with white, with
a fine white basal line and three black lines alternating with white.
Hind wing dirty whitish, touched with grey towards apex, veins slightly darker
ochreous, cubital pecten grey. Cilia pale fuscous with a darker fuscous subbasal
shade.
Male genitalia. Tegumen high. Uncus moderately long, slender. Socius moderate.
Vinculum strong, flattened, smooth. Valva long; cucullus with a slender top and
base, dilated in middle, costa prominent above middle. Valvula with a long, swol-
len stalk which has a rounded prominence in middle, and a series of moderate
spines, decreasing in size basally; top of valvula moderately broad; sacculus strongly
dilated and flattened, semicircular, moderately concave, simple; a small patch of
punctulations above centre and a dense corona of long and slender spines along
outer edge. Juxta small and strong. Aedeagus very long, hooked downward,
sclerotized.
® 13—19 mm (allotype, 16 mm). Head, palpus, and tegula as in male, thorax
tawny. Abdomen ochreous-fuscous.
Fore wing slightly narrower and longer, otherwise of the same shape, costa
slightly less curved, apex and termen similar. Markings of the allotype which is
slightly rubbed, are similar, except that the ground colour appears paler, whitish,
A. DIAKONOFF: African species of Bactra 339
perhaps due to rubbing, while there is never a trace of the ocellus nor of its central
dark dot.
Hind wing similar to that in male, apex variably suffused with greyish.
Sometimes the entire wing in the female suffused with bright ochreous-tawny,
apical streak deep brown, connected with base of wing by a suffused fuscous streak,
containing a dark fuscous first discal spot; stigma conspicuous (e.g., © paratypes
nos. 3794, Karkloof, and 3646, Kimbolton).
Female genitalia. Lobus analis triangular, pointed, inner edge below middle with
an excision. Postapophyses long and slender, anapophyses considerably shorter.
Sterigma moderately sclerotized mesially, margin of sclerotization ill-defined towards
rostral edge which is triangular-truncate; above this edge a gradually curved strong
tubular fold with irregularly denticulate structure. Ostium shaped as a triangular
funnel, sclerotized; colliculum, a strong tube, beginning immediately below the
edge of the rather small lamella antevaginalis, calyciform above, with thin wall,
cylindrical below, with thicker wall. Cestum present, a small triangular sclerite.
Corpus bursae normal. Signum basket-shaped, rather shallow.
Material studied. South Africa, Transvaal: Pretoria, 30.1.1910, holotype, &,
gen. no. 4295; allotype, @, 31.1.1910, gen. 4273. Paratypes, the same locality,
291007 290478) 25-111.1910, "gen, no, 3459782 (British Museum);
27.1.1910, (4284 3); 29.11.1909 (4274 4); 8.1.1910 (4239 3); 12.11.1910
(4276 &); the same locality, 28.11.1910 (4275 2); 20.III.1909 (4288 9)
(A. J. T. JANsE). Nelspruit, 1 ¢, XII.1917 (4238) (H. G. Breijer). Natal:
Weenen, Kimbolton Estcourt, 1892, 1 4, gen. no. 3640; 2.9 3645 and 3646
(HITCHINSON) (British Museum). Nkwaleni, 2 &, 10.1.1916, 4285 and 12.I.-
1916, (4272); 2 9, 10.1.1916, 4268 and 4269 (A. J. T. JANSE); Magude,
IV.1918 4293 & (C. J. SWIERSTRA). Karkloof, 11.1.1917, 1 9, 3794 (A. J. T.
JANSE). Indaleni, 11.1951 1 9, 4231 (J. W. Hunt). Cape Province, Umtata,
2 OprmilestE SSA IA 93 ieal wa 705 MO 379922] Le ANSE) 2 Instotal 12 07;
10) 79%
Allied with B. (C.) coronata Diak., from Java, Borneo and the Philippine
Islands.
Bactra (Chiloides) sinassula spec. nov.
Figs. 50—52
& 19—21 mm (holotype 18 mm). Head and thorax tawny-fuscous; palpus
rather long, abruptly dilated; tawny, infuscated. Abdomen fuscous.
Fore wing oblong-subtriangular, costa gently curved at extremities, apex pointed,
termen clearly sinuate, oblique. Rather pale glossy purplish-fuscous. Markings
fine, dark fuscous, partly edged or suffused with tawny-olive. Costal markings
narrow, dark, along anterior half short, along posterior very slender, termen from
apex to dorsum with a dark fuscous line; first discal spot indistinct, represented
by a whole series of transverse fuscous marks more or less interconnected so as to
form zigzag marks, second discal spot obliterate; all veins on posterior half of
wing very finely streaked with dark fuscous; upper half of posterior portion of
cell rather distinctly filled out with brighter tawny-ochreous; entire wing with
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
340
A. DIAKONOFF : African species of Bactra 341
minute transverse strigulae between the veins. Cilia (imperfect) concolorous, with
an antemedian fuscous line.
Hind wing pale fuscous becoming whitish towards dorsum; deeper fuscous
towards apex. Cilia glossy whitish-fuscous.
Male genitalia. Tegumen and vinculum very broad, the latter smooth. Valva
with cucullus moderately broad, sinuate, costa strongly prominent and rounded
beyond base, top of cucullus gradually narrowed, spines moderate. Valvula rather
long with blown up stalk, bearing some five spines below the corona which
gradually become longer. Sacculus extremely large, over 1/,, dilated and extended,
lower edge gradually rounded, its outer half with a single row of very long,
slender spines; punctulate area small, at the base of valvula. Juxta very broad.
Caulis moderate. Aedeagus long, sclerotized, bent downward.
The series is rather variable. One paratype (4482) is paler, suffused with tawny-
olive and tawny-ochreous, with a whitish-ochreous faint supramedian streak from
base to apex, along fold edged below with cloudy dark fuscous suffusion. Two
males (3802, 3803) are melanistic, more or less densely infuscated throughout.
Finally the male no. 4252 is similar to the type but has a distinct second discal
spot and indications of a pale stigma.
Material studied. South Africa, Transvaal: Pretoria, 15.XII.1911 (A. J. T.
JANSE), holotype, 3, gen. no. 4472. The same locality and collector, 1.1918, 1 4,
paratype, no. 3602; the same, 16.11.1909, 1 &, paratype, gen. no. 3803. Natal:
Magude, IX.1918 (C. J. SWIERSTRA), 1 &, paratype, gen. no. 4399; the same
locality and collector, X.1918, 1 3, paratype, gen. no. 4252. Also one &, para-
type: British East Africa (F. THOMAS), gen. no. 4482 (in the Vienna Museum).
Gee
9 16—20 mm (allotype 19 mm). Head and thorax in middle creamy, laterally
and on tegulae suffused with very pale fulvous-tawny. Palpus pale tawny with a
whitish base and an oblique median dark grey shade. Abdomen fuscous.
Fore wing oblong, rather narrow, costa gently curved at base, apex pointed,
termen sinuate, oblique. Pale ochreous, markings tawny, fuscous, and dark brown.
Costa with a series of fine and short dark brown strigulae. An ill-defined longitu-
dinal median suffusion, greyish-fuscous mixed with tawny and dark brown,
extending to above dorsum; on anterior half this suffusion cut by several brown
ill-defined inwards-oblique marks, anterior three of these representing the first
discal spot; second discal spot shaped as some faint dark edging to lower half of
stigma, which is white and little contrasting; the median streak considerably dilated
and brighter tawny-brown beyond the cell; a pale creamy streak above cell from its
middle to apex; veins finely streaked with brown; termen with a dark brown line;
dorsum with a series of dark brown dots. Cilia tawny, strongly dusted with creamy,
a subapical and an apical dark brown lines.
Hind wing and cilia glossy, creamy-white.
The series is rather variable, one specimen brighter pale ochreous with more
Fig. 51. Bactra (C.) sinassula sp.n., 9, allotype, sterigma and ovipositor. Fig. 52. The same,
bursa. Fig. 53. B. (C.) salpictris sp.n., &, holotype, sterigma and ovipositor. Fig. 54.
The same, bursa
342 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
extended costal strigulation and median streak brighter tawny ochreous; two other
specimens more greyish-fuscous tinged, with a darker but interrupted median
streak, containing one or both discal spots.
Female genitalia. Sterigma sclerotized, strongly folded at the sides, two longer
folds flanking the osttum bursae which is calyciform and wide, with a pair of
short parallel folds on each side above extensions of the edge of ostium. Col-
liculum, a very strong, straight tube, gently dilated above, with strongly thickened
walls below dilatation, and complicated structures of the inner surface. Anapophy-
ses are short, with peculiar rectangularly bent bases. On each side of the colliculum
there is an oval tumescence. Postapophyses very long and slender. Signum large,
basket-shaped.
Material studied. South Africa, Transvaal: Pretoria, 13.1.1916 (A. J. T. JANSE),
1 9, allotype, gen. no. 4270. Natal: Bela Vista, XI.1916 (C. J. SWIERSTRA),
1 @, paratype, gen. no. 4236; Tanganyika: Victoria Falls, 26.1.1918 (Coll. JANSE),
1 9, paratype, gen. no. 4291.
Comoro Ids.: Mayotte (L.), V.1911 (in MEyYRICK’s hand), 1 & paratype, gen.
no. 3600 (in MEYRICK Collection, in the British Museum).
It seems very likely that the sex partners are correctly chosen because the long
and sclerotized aedeagus corresponds with the long colliculum.
A large and robust species, belonging to the coronata group.
Bactra (Chiloides) triceps spec. nov.
Figs. 58—60
& 15—17 mm (holotype 15 mm). Head sordid whitish, tinged creamy, tuft
on vertex, which is rough and high, mixed with fuscous-tawny laterally. Palpus
strongly dilated, with rough edges, fuscous-grey externally, creamy inside, terminal
segment very slender, grey. Thorax pale ochreous, regularly spotted with fuscous,
metathorax with a silvery gloss. Abdomen fuscous.
Fore wing oblong, little dilated, moderately broad, costa gently curved, more
so along posterior half, apex rather obtuse, termen gently convex, almost straight,
little oblique. Ochreous-white, with a faint gloss, partially clouded with pale
fuscous-grey. Markings dull, ferruginous and dark fuscous. A conspicuous streak
of pale ground colour from beyond base to above apex, rather well-defined, oc-
cupying about 1/, of wing breadth, gradually attenuated at extremities, running
along upper edge of cell; remainder of wing clouded; costa along posterior part
with distinct ferruginous little oblique strigulae, delimited by the white streak,
along anterior part of costa obliterated; median longitudinal fascia irregular but
complete, from base to apex, dark fuscous, obscuring discal markings; first discal
spot discernible, second discal ill-defined, followed by two roundish dark brown
suffused spots between cell and termen; an oblique-oval moderate spot forming
the centre of ocellus; a narrow slightly undulate or dentate terminal streak, above
connected by a curved slender line with costa. Cilia grey, with several faint darker
lines, in tornus pale ochreous-grey, along costa pale ochreous.
Hind wing sordid whitish-ochreous, with a moderate gloss, slightly infuscated
towards apex. Cilia pale ochreous, with a golden gloss around apex.
A. DIAKONOFF : African species of Bactra 343
%
Li
dI
su
fas
D:
Pat >
SEO
Fig. 55. Bactra (C.) trimera sp.n., 6, holotype, genitalia. Fig. 56. The same, ®, allotype,
sterigma and ovipositor. Fig. 57. The same, bursa
344 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
Male genitalia. Very characteristic. Tegumen short and broad, vinculum widely
V-shaped. Uncus rather short. Socius large. Valva peculiar, cucullus narrowed at
base, costa rounded and strongly prominent below middle, top considerably at-
tenuated, slender. Valvula moderate, with a large corona. Sacculus strongly concave,
top extended so as to form a large triangular process, with 7—8 long spines along
edge; base of sacculus narrowed, concave; punctulated area on an oblique transverse
discal tumescence. Juxta large. Caulis short. Aedeagus robust, rather short, cornuti,
some 5 long spines.
Material studied. South Africa, Natal: Bela Vista, XI.1914 (C. J. SWIERSTRA),
1 4, holotype, gen. no. 4243; Magude, IX.1918 (C. J. SWIERSTRA), 1 4, para-
type, no. 4241. Transvaal: Woodbush Village, 1670 m, 11.1.1925 (A. J. T.
JANSE), 1 4, paratype, no. 4367.3 3.
The male from Woodbush is melanistic, with the fore wing strongly infuscated,
but with discal markings, the discal spots, and the pretornal dot remaining distinct.
The second paratype is similar to the holotype.
9 18—24 mm (allotype 22 mm). Head and thorax tawny-fulvous. Palpus pale
fulvous, basal half of tuft of median segment blackish. Abdomen fuscous.
Fore wing oblong, rather narrow, costa curved towards extremities, straight in
middle. Pale fuscous, glossy, evenly suffused except above tornus, with dull tawny.
Costal markings reduced to minute dark purplish traces along extreme edge
posteriorly; first and second discal spots indicated by some dark purple dustings;
traces of some leaden transverse strigulae below costa posteriorly; termen with a
tawny line dusted with purplish; part of wing below vein 7 and as far as cell
rather well-defined and not suffused with tawny. Cilia purple, dull, dusted with
whitish, a narrow basal and an indistinct subapical whitish lines.
Hind wing glossy, slightly suffused with grey. Cilia concolorous.
A variable lot. Except the allotype there are three tawny-greyish unicolorous
specimens, two pale ochreous insects with a narrow grey median streak, faint and
interrupted, and two pale ochreous specimens with well-defined dark fuscous-grey
broad median streak, in one of them gradually dilated so as to occupy almost
entire termen.
Female genitalia. Seventh sternite strongly sclerotized and plicate, outer rostro-
lateral margin stronger sclerotized and with a narrow dark fold on each side.
Ostium rather wide, forming a strong funnel, its walls thickened along upper
portion from below top. Anapophyses rather long and slender, postapophyses
slender and long. Signum, a shallow basket.
Material studied. In the Transvaal Museum. South Africa, Natal: Karkloof,
23.1.1917 (A. J. T. JANSE), 1 2, allotype, gen. no. 4479. All other are 9
paratypes: Transvaal: Tweefontein, 13.1.1907, gen. no. 4299; Pretoria, 7.1.1910,
gen. no. 2879; 5.XII.1907, gen. no. 4354; the same, 15.11.1915, gen. no. 4302;
the same, 6.XII.1917, gen. no. 4304. Barberton, 31.XII.1910, gen. no. 4307 (all
by A. J. T. JANSE).
In the British Museum. Natal: Weenen, X-XI.1925 (H. P. THOMASSET), 1 2,
paratype, gen. no. 3627. Transvaal: Pretoria, 2.XI.1907 (JANSE), gen. no. 3462
(MEYRICK Collection). 8 9.
A. DIAKONOFF : African species of Bactra 345
Bactra (Chiloides) nesiotis spec. nov.
Figs. 61—62
@ 19 mm. Head and thorax dark fuscous-grey. Palpus rather pointed, pale
ochreous-tawny, with a well-defined median blackish line running to tip. Abdomen
rather dark fuscous-grey.
Fore wing narrow, sublanceolate, costa rather curved along anterior half, almost
straight posteriorly, apex acutely pointed, termen almost straight, gently convex,
very oblique. Rather unicolorous dark fuscous-grey. Costa slightly lighter, slaty-grey
with minute linear transverse black marks; basal third of wing tinged purplish-
grey, darker; terminal portion of wing somewhat more tinged grey-olive, dorsum
with a series of approximated triangular dark fuscous dots. Cilia sordid pale
ochreous-fuscous mixed with fuscous, with slightly paler base and ill-defined sub-
basal line.
Hind wing dark fuscous with a bronze gloss, paler towards costa; in paratype
hind wing dull fuscous-black, unicolorous. Cilia pale fuscous with a fuscous sub-
basal band.
Female genitalia. Lobi anales slender and elongate. Sterigma formed by a
transversely oblong moderately sclerotized lamella postvaginalis, its upper edge
with two triangular excisions, middle part rounded. Lower edge of this sclerite
forming the ostium bursae, continued as a strong tubular colliculum, gradually
dilated downwards, with a split in middle of lower third. Ostium flanked by
slightly sclerotized plates, finely wrinkled along inner edge which is prolonged
downwards. Signum basket-shaped.
Material studied. Madagascar: Antananarivo”, (M.), 1911, 1 ®, holotype
(rather worn, right fore wing and head missing), gen. no. 2158. Paratype, 1 9,
the same data, without abdomen. (Labels in MEYRICK's hand. In MEYRICK
Collection, in the British Museum).
A conspicuously dark insect with strongly pointed fore wings. Superficially it
resembles a Nannobactra and is, in fact, very similar to B. (Nannobactra) sardonia
Meyr. Its position is obscure without the evidence of the male genitalia.
The genitalia resemble slightly those of contraria Diak. from Borneo, but resem-
blance may be purely incidental.
Subgenus Nannobactra Diakonoff, 1956
Nannobactra Diakonoff, 1956, Zool. Verh., no. 29, p. 52 (subgen. of Bactra).
The subgenus differs from Chiloides by compact male genitalia with the valvula
present and equally well-developed, but with the cucullus distinctly separated
from the basal part of the valva by a transverse groove; this cucullus is usually
rounded, sclerotized throughout and beset along its outer edge with one to three
series of very large spines becoming smaller towards top of cucullus. The females
possess a weakly sclerotized sterigma, formed by little modified, small and usually
finely short-haired ridges, tumescences and folds. The signum is a small concave
and denticulate sclerite, often weak, sometimes absent.
The species usually are small, with narrow and pointed wings, and with similar,
346 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
Fig. 58. Bactra (C.) triceps sp.n., 6, holotype, genitalia. Fig. 59. The same, 9, allotype,
sterigma and ovipositor. Fig. 60. The same, bursa
although variable markings. It is very easy to identify the subgenus, but the species
usually are difficult of recogniton, especially the females. Although I am including
the males in the key to the species, I am not able to do the same for the females,
except for the very distinct sardonia. It is likely that only rearing of whole egg
A. DIAKONOFF : African species of Bactra 347
batches will allow certain identification and separation of the females of Nanno-
bactra species; a key for that purpose is not possible at present.
To stress and elaborate certain differences of the structure of the male genitalia
in this difficult group, I am introducing a few new (non-Latin) descriptive terms.
The degree of the development of the gnathos, or at least, of structures which
seem to be homologous with that part, is of specific importance. Sometimes a
membraneous, complete transverse and band-like gnathos is present, more often
it is obliterated in the middle and only its basal parts, shaped as triangular lobes
at the tegumen, are visible; for these structures I use the therm ’’shoulder lobes”.
Sometimes they may be entirely absent. The large spines along the edge of the
cucullus, are homologous with ’’cucullus bristles and spines’, an indication used
by me before, in the subgenera Bactra and Chiloides. In Nannobactra these spines
are so large that they form a marginal corona; therefore the term “corona spines”
is used for these structures. Often the ultimate and the penultimate 1—3 corona
spines are much larger or differently shaped than the other; in this case they are
termed ’’inequal’’. Seldom this difference is slight, then they are termed "subequal
corona spines”.
Bactra (Nannobactra) sardonia (Meyrick, 1908)
Figs. 64, 67—68
Pamplusia sardonia Meyrick, 1908, Proc. Zool. Soc. Lond., p. 718 (4 9). — Clarke, 1955,
Meyrick’s Types, vol. 1, p. 277.
Bactra sardonia: Clarke, 1958, loc. cit, vol. 3, p. 315, t. 156 figs. 1—1a (lectotype
select., fig.).
Distribution. Transvaal: N.E. Pretoria.
This apparently rare species has hitherto been known after the unique male
lectotype, in the British Museum. By a lucky chance a single female specimen, in
my opinion undoubtedly conspecific with the male, was found in the MEYRICK
Collection. This female and the genitalia of the two sexes may be described as
follows.
‘9 17 mm. Head and thorax pinkish-fulvous. Palpus whitish, upper edge and
tip fulvous. Abdomen dark fuscous.
Fore wing sublanceolate, little dilated, costa gently sinuate, being curved along
anterior half, slightly prominent before middle and hardly concave posteriorly,
apex sharply pointed, termen gently concave, almost straight, very oblique. Evenly
tawny-fulvous, only towards costa becoming paler, fulvous-greyish. Costa with
numerous dark brown, small, wedge-shaped transverse marks, on posterior half of
costa with whitish interspaces, which become pale pinkish before apex. Cilia
pinkish-fulvous, mixed with whitish, with a pure white basal streak, a cloudy black
bar opposite apex.
Hind wing dull dark bronze-fuscous, darker towards apex, paler towards dorsum.
Cilia (imperfect) fuscous with a dark fuscous subbasal band.
Male genitalia. Tegumen broad, spherical. Uncus large; socius small. Gnathos
indicated, very weak. Valva of the usual shape, but valvula very large and
characteristic, corona being continued along the ventral edge of the valvula to its
base. Sacculus small, with a large punctulate area, spined only in centre and to-
348 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
A. DIAKONOFF : African species of Bactra 349
wards the outer edge, spines moderate, not numerous. Aedeagus rather long.
Female genitalia. Eighth + ninth segment sclerotized, intersegmental membrane
between this and lobi anales finely aciculate ventrally, naked laterally. Both
apophyses pairs quite long. Sterigma well-modified, lamella posterior forming a
rather narrow vertical prominence, densely aciculate, with a truncate top. Ostium
bursae pear-shaped, its lower edge gradually rounded and thickened; lamella
antevaginalis developed, a sinuate transverse band or fold with thickened edges,
twice sinuate; curved downwards in the middle and again, less so, at the sides;
strong folds running from the extremities of this lamella along edges of the genital
segment, and becoming weaker and less sclerotized dorsad. Signum, a small concave
sclerite, finely scobinate and denticulate.
Material studied. South Africa, Transvaal: N.E. Pretoria, 14.1.1907 (JANSE),
®, gen. no. 3519. (In MEYRICK's hand; in MEyRIcK Collection, in the British
Museum).
Bactra (Nannobactra) legitima Meyrick, 1911
Figs. 68, 70—71
Bactra legitima Meyrick, 1911, Trans. Linn. Soc. Lond., vol. 14, p. 269 (2). — Viette,
1947, Mém. Inst. sci. Madagascar, ser. A. vol. 1, p. 54. — Clarke, 1955, Meyrick’s Types,
vol. 1, p. 183. — 1958, loc. cit., vol. 3, p. 312, t. 155 fig. 1 (type figured).
Bactra (Nannobactra) legitima: Diakonoff, 1961, Ann. Soc. ent. France, vol. 130, p. 70
(Mahé, 4 neallotype).
Distribution. Seychelles: Silhuette Id, Mahé Id.
Material studied. South Africa, Natal: St. Lucia Lake, X.1920 (H. W. B.),
DS cen) no 4379-5 New = Hanover,” 26X11 1915, CA ]. T. JANSE), 378,
gen. nos. 3792, 4335, and 4339. Southern Rhodesia: Salisbury, 1.1.1920 (A. J. T.
JANSE), 1 4, gen. no. 4543 TM. Bulawayo, 15-23.X11.1919 (A. J. T. JANSE),
ES, gen. no. 4370.
Besides, a 9, with a label in MEYRICK’s hand: "Silhouette, Seychelles I., X.09”
(Meyrick Collection, in the British Museum), gen. no. 2157. A worn specimen
without head, but with an intact abdomen, obviously conspecific with the female
type specimen (without abdomen) in the British Museum and, very probably,
being the paratype. Female genitalia may be described as follows.
The genital segment moderately and evenly sclerotized throughout; this segment
discending on each side of sterigma, so as to form slender smooth bands, each
ending in a small free lobe level with the ostium; these bands sharply delimiting
the sterigma proper which is weak and not sclerotized but recognisable by fine
aciculation (while the bands are naked); lamella postvaginalis with a curved upper
edge; ostium wide and regular; colliculum well-defined but neither sclerotized, nor
aciculate and rather shallow.
Bactra (Nannobactra) minima Meyrick, 1909
Bactra minima Meyrick, 1909, Journ. Bombay Nat. Hist. Soc., vol. 19, p. 586 (4 Bar-
beryn Island, Ceylon). — Diakonoff, 1950, Bull. Brit. Mus., Ent., vol. 1, p. 288, pl. 6
Fig. 61. Bactra (C.) nesiotis sp.n., 9, holotype, sterigma. Fig. 62. The same, signum.
Fig. 63. Lobesia (Lomaschiza) quadratica (Meyr.), ®, holotype, genitalia. Fig. 64. Bactra
(Nannobactra) sardonia Meyr., 8, holotype, genitalia. Fig. 65. Lobesia (Harmosma
harmonia (Meyr)., neuration of right fore wing
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
A. DIAKONOFF : African species of Bactra 351
fig. 25 (lectotype designated, gen. 4 descr. and fig.). — 1956, Zool. Verh., no. 29, p. 55,
figs. 52—53 (phaeopis syn.).
Bactra phaeopis Meyrick, 1911, Proc. Linn. Soc. N.S. Wales, vol. 36 p. 254 (4 2, Sudest
Island, New Guinea). — Diakonoff, 1950, Bull. Brit. Mus., Entom., vol. 1, p. 288, pl. 6
fig. 28 (gen. 4), pl. 8 fig. 41 (gen. &) (Lectotype designated, gen. &, 9 descr. and
fig.).
Distribution. Ceylon (Barberyn Island); New Guinea (Sudest Island); Solomon Islands;
Marianas Islands.
Canary Islands, Tenerife, IV.1884 (LEECH) (WALSINGHAM Collection ), gen.
no. 3697, & (no. 62294), 3700 & (78866), 3701 & (78865) and 3705 9
(78864). All in WALSINGHAM Collection, British Museum.
It is surprising to find minima "so far from home”. This species must be widely
distributed, but must have escaped attention of collectors due to its modest
colouring and size.
The male genitalia of the present material agree in all details with those of the
Asiatic specimens. The female has an exactly similar sterigma, but the genital seg-
ment is somewhat more sclerotized. However, this feature is subject to some
variation throughout the subgenus.
Bactra (Nannobactra) dasioma spec. nov.
Bigs 6947/3
4 16 mm. Head creamy, vertex suffused with pale tawny. Palpus whitish above,
creamy, mixed with light brown, laterally. Thorax creamy, spotted with fuscous.
Abdomen light grey, anal tuft pale ochreous.
Fore wing rather broad, oblong, costa curved at extremities, straight in middle,
apex moderately pointed, termen straight, oblique. Creamy-white, glossy. Markings
light tawny-brown, fuscous and blackish, well-defined. Costa along anterior half
with not numerous dark fuscous wedge-shaped small marks, posterior half of costa
contrastingly different: with six broad, light tawny transverse spots; first of these
median and trapezoidal; second narrow, fasciate; third to fifth, wedge-shaped;
sixth apical, rounded; spaces between these spots formed by glossy pairs of white
costal marks, each pair with a minute separating median dark line; fourth costal
mark continued by a straight streak to upper part of termen; first discal spot
arrowhead-shaped, black, continued below as an inwards-oblique streak across wing
to above dorsum; second discal spot semioval, longitudinal, dark tawny-fuscous,
on middle of its upper edge a small tawny suffusion indicating rest of transverse
fascia; stigma small and pale, upper half absent; a slightly inwards-oblique short
fuscous streak between cell and tornus; a pair of irregular curved ochreous-tawny
marks on upper part of termen, united into a suffused triangular fuscous marginal
line from apex to tornus. Cilia creamy, mixed with pale ochreous, a pale ochreous
antemedian line, a dark fuscous small blotch opposite apex.
Hind wing glossy pale ochreous, considerably suffused with light purplish, more
so towards apex.
Fig. 66. Bactra (N.) legitima Meyr., 4, neallotype (after Diakonoff, 1961). Fig. 67.
| B. (N.) sardonia Meyr., 9, gen. no. 3519. Fig. 68. The same, signum. Fig. 69. B. (N.)
dasioma sp.n., ©, allotype, sterigma and ovipositor, Fig. 70. B. (N.) legitima Meyr., ©,
gen. no. 2157. Fig. 71. The same, signum
292 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
9 16 mm. Exactly similar to the male but with all markings somewhat darker,
fuscous, costal spots narrower, pretornal mark broader, touching dorsum before
tornus, first discal spot continued to dorsum. Hind wing suffused with darker
purple, except towards base.
Male genitalia. Comparable with those of phaulopa Meyrick from Asia, but
clearly differing by the presence of shoulder lobes which are slightly rounded
and aciculate. Valva of normal shape, corona spines large and long, in two oblique
series, punctulate area with only few thick and rather short spines (10 left, 11
right), and a few fine teeth on the inner edge; lower part of sacculus extended
into a broad blade, rounded below, forming a short triangular cusp above.
Female genitalia of phaulopa type, but more similar to those in verutana Zell.
The genital segment sclerotized, lamella antevaginalis with upper and lower edges
distinct and sclerotized. Small lateral sclerites present, separate and very dark.
Signum minute.
South Africa, Cape Province: Stella Bush, 28.11.1915 (A. J. T. JANSE), 1 4,
holotype, gen. no. 4477. Natal: Umkomaas, 29.1.1914 (A. J. T. JANSE), 1 9,
allotype, gen. no. 4459. (In the Transvaal Museum).
Cape Province: Umtata, X.1904 (LEIGH), 16411, 1 &, paratype, gen. no.
3655 (WALSINGHAM Collection, in the British Museum). 2 &, 1 2. A brightly
marked species with, for the subgenus Nannobactra, unusually broad fore wings.
Bactra (Nannobactra) aciculata spec. nov.
Fig. 72
& 14 mm. Head, palpus and thorax whitish, shoulder fuscous. Abdomen
fuscous.
Fore wing sublanceolate, dilated, broadest at 3/4, costa gently curved troughout,
more so along its posterior fourth, apex acutely pointed, termen tolerably straight,
oblique. Creamy-white with a silky gloss, markings dark fuscous (rather worn).
Anterior part of costa narrowly suffused with dark grey and with a series of dark
fuscous transverse marks; basal patch not defined, transverse fascia indicated by
a moderate transverse fuscous spot on middle of costa, below costa extended into
a fasciate longitudinal small tawny suffusion, continued across wing by a very
faint pale yellowish colour; first discal spot longitudinal, fasciate; second discal
spot irregularly semioval, rather well-defined, dark fuscous, its ends acutely produc-
ed, stigma represented by a small excision of its upper edge; apical streak faint
anteriorly, posteriorly blotchy and emitting a vertical bar to lower part of termen;
a jet-black oval spot in apex; faint transverse marks along dorsum. Cilia dark
fuscous (damaged).
Hind wing and cilia pale fuscous (imperfect).
Male genitalia of phaulopa type. There is a distinct membraneous and band-like
gnathos. Cucullus with a broadly rounded top, corona spines in two series,
moderately curved and subequal (ultimate 3—4 spines hardly different in shape
and size), pointed. Sacculus as large as cucullus, lower edge with a moderate,
gradually rounded blade, becoming narrower above, cusp blunt; punctulate area
with a moderate number of strong pointed spines, situated towards top and outer
edge; inner edge of this area with several inequal, very acute teeth.
A. DIAKONOFF : African species of Bactra 353
Fig. 72. Bactra (N.) aciculata sp.n., &, holotype, genitalia. Fig. 73. The same of B. (N.)
diasoma sp.n., & holotype
Distinct by strongly rounded cucullus, subequal corona spines, position of
sacculus spines and dentate inner edge of punctulate area.
Material studied. South Africa, Southern Rhodesia: Selukwe, 27.X11.1919 (A.
J. T. JANSE), 1 4, holotype, gen. no. 4466. Unfortunately a single specimen
available; the species, however, is sufficiently distinct.
354 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
Bactra (Nannobactra) spec.
The following six female specimens of Nannobactra must remain unidentified
for the present, due to their poor condition and to the considerable variability of
the sclerotization of the genital segment in the species of this subgenus.
3788, Mazoo (S. R.), 1.1920 (Coll. JANSE). 4329 and 4330, Barberton (T.),
15.1.1911 and 31.XII.1910 (A. J. T. JANSE). 4335, Durban (C. P.), 13.XII.1916
(v. D. MERWE, Coll. JANSE). 4369, Bulawayo (S. R.), 15-23.XII.1919 (A. J. T.
JANSE), and 4402, Lorenzo Marques, 23.11.1930 (A. C. DAINTREE).
APOCRYPHAL SOUTH AFRICAN Bactra SPECIES
Two following species have been originally described as, or subsequently
transferred to, Bactra.
Lobesia Guenée, 1845
Harmosma subgen. nov.
Fig. 65
With superfical characters of Lobesia but with veins 9 and 10 moderately
sinuate, approximated posteriorly, then slightly diverging again; pterostigma ill-
defined.
Male genitalia with tegumen weak, rounded. Gnathos membraneous, a weak
curved transverse rod. Tuba analis ill-defined. Valva rather narrow, sclerotized
throughout, cucullus slender, narrowed, covered all over, but not densely, with
slender spines; sacculus strongly thickened, with a transverse deep groove (scallop-
ed), separating Spc, and Spc, completely; both these spine clusters large, elongate,
formed of extremely dense spines, those on the ventral end of Spc, very long,
forming a projecting sheaf.
Female genitalia with sterigma shaped as a long, entirely free, rising tube,
narrowed towards the top which is spinulose. Signum, a small denticulate concave
sclerite.
Type-species, Polychrosis harmonia Meyrick, 1908. An interesting form, deriv-
able from the Palaearctic subgenus Lobesiodes Diakonoff, 1954.
Lobesia (Harmosma) harmonia (Meyrick, 1908) combin. nov.
Fig. 65
Polychrosis harmonia Meyrick, 1908, Proc. Zool. Soc. Lond., p. 716 (4 9, Transvaal:
Pretoria). — 1911, Ann. Transv. Mus., vol. 2, p. 63 (transferred to Bactra). — Clarke
1955, Meyrick’s Types, vol. 1, p. 153.
Bactra harmonia: Clarke, 1958, Meyrick’s Types, vol. 3, p. 311, t. 154 figs. 2—2a (lecto-
type selected, figured).
Distribution. South Africa; Madagascar.
The genitalia, as described for the subgenus above.
A. DIAKONOFF : African species of Bactra 355
Lobesia (Lomaschiza) quadratica (Meyrick, 1912) combin. nov.
Fig. 63
Bactra quadratica Meyrick, 1912, Ann. Transv. Mus., vol. 3, p. 63 (9, Natal: Nkwaleni).
— Clarke, 1955, Meyrick’s Types, vol. 1, p. 267. — Anonymous, 1958, List Types S. & E.
Afr. Museums, vol. 1, p. 57 (holotpe indicated).
Distribution. South Africa.
APPENDIX
After the manuscript of the present paper has been sent to the printer, I was
entrusted with a small collection of Bactra species, collected in Sudan, which were
kindly sent to me for identification by Dr. KLAUS SATTLER of the Munich Mu-
seum, Germany. It seems worth while to add these latest data to my paper, for the
sake of completeness.
Bactra (Bactra) graminivora Meyrick, 1922
This paper, p. 312
Sudan, Ed Damar, Hudeiba, 1.X.—18.XI.1961 (R. REMANE), 2 4, 14 9,
genitalia nos. & 4615, 4622; 9 4604—4608, 4610—4614, 4616—4618 and
4621.
Bactra (Chiloides) venosana (Zeller, 1847)
This paper, p. 331
Sudan, Ed Damar, Hudeiba, 18.X1.1961, 15.11.1962; Blue Nile Province, Wad
Medani, 3.VIII.1962 (R. REMANE), 3 6,1 9, genitalia nos. 4542 4, 4620 4,
4609 9.
Bactra (Chiloides) tornastis Meyrick, 1909
Bactra tornastis Meyrick, 1909, Journ. Bombay Nat. Hist. Soc., vol. 19, p. 586 (4 9
Ceylon; Coorg). — Diakonoff, 1950, Bull. Brit. Mus., Ent., vol. 1, p. 289, t. 5 fig. 18
(lectotype select., genit. figured). — Clarke, 1955, Meyrick’s Types, vol. 1, p. 312. —
1958, loc. cit., vol. 3, p. 315, t. 156 figs. 3—3a (lectotype illustr.).
Distribution. India: North Coorg; Ceylon; Pakistan.
Sudan, El Damar, Hudeiba, 18.X1.1961, gen. no. 4619 3. Blue Nile Province,
Wad Medani, 3.VIII.1962, gen. no. 4624 ¢ (R. REMANE). 2 ¢.
The discovery of this Indian species in Africa is a complete surprise. The species
is intermediate between fasciata and triceps, but all three are entirely distinct. In
my key tornastis would run to no. 14, triceps. However, the male genitalia differ
by the sacculus being less extended laterad, having a more rounded and less promi-
nent point, beset with a much longer series of distinctly shorter spines. The genita-
lia agree with those of my Asiatic examples of tornastis closely. There is no doubt
of this being the same species.
356 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 7, 1963
REFERENCES
DIAKONOFF, A., 1950. — The type specimens of certain Oriental Eucosmidae and Carposin-
idae (Microlepidoptera) described by Edward Meyrick, together with descriptions
of new Eucosmidae and Carposinidae in the British Museum (Natural History).
Bull. Brit. Mus., Ent., vol. 1, p. 285—289, t. 5—8.
, 1956. — Records and descriptions of Microlepidoptera (8). Zool. Verh., no. 29,
p. 1—60, figs. 1—57.
———, 1959. — Further records and descriptions of Bactra species (Lepidoptera, Tortric-
idae), chiefly in Dr. H. G. Amsel Collection. Bijdr. Dierk., part 29, p. 173—186,
figs. 1—18, table.
— 2, 1962. — Preliminary survey of the Palaearctic species of the subgenus Bactra
Stephens (Bactra, Tortricidae, Lepidoptera). Zool. Verh., no. 59, p. 1—48, figs.
1—36, t. I—X VIII.
MABILLE, P., 1900. — Lepidoptera nova malgassica et africana. Ann. Soc. ent. France, vol.
68 (1899), p. 751.
Meyrick, E., 1909. — Descriptions of Transvaal Microlepidoptera. Ann. Transv. Mus.,
Vola 2Np Sta SNEL
, 1911. — Descriptions of Transvaal Microlepidoptera, II and III. Ibid., vol. 4,
p. 62—63, 227—228.
WALSINGHAM, Lord, 1881. — On the Tortricidae, Tineidae and Pterophoridae of South
Africa. Trans. ent. Soc. Lond., 1881, p. 231—232.
WALKER, F., 1866. — List Lep. Het. Brit. Mus., vol. 35, p. 1750.
ZELLER, P. C., 1852. — Lepidoptera microptera quae J. A. Wahlberg in Caffrorum terra
collegit, p. 84, Stockholm.
Index to species
Synonyms are in italics
aciculata 352
adelpha 316
jansei 313
lactosana 318
scrupulosa 300
siccella 326
aletha 334
Bactra, subgen. 294
Chiloides, subgen. 313
confusa 308
crithopa 312
dasioma 351
dolia 300
endea 321
fasciata 338
furfurana 310
graminivora 312, 355
harmonia 354
Harmosma, subgen. 354
hebetata 296, 298
lanceolana 310
legitima 349
Lobesia 354
Lomaschiza 355
minima 349
Nannobactra, subgen. 345
nesiotis 345
pauperrima 318
punctistrigana 330
pythonia 296
quadratica 355
rhabdonoma 322
salpictris 336
sardonia 347
simpliciana 318
sinassula 339
sordidata 318
Spinobactra, subgen. 291
spinosa 291
stagnicolana 326
tornastis 355
tradens 305
triceps 342
trimera 331
tylophora 319
venosana 331, 355
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DEEL 106 AFLEVERING 8 2 1963
TIJDSCHRIFT
VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
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INHOUD:
B. M. LENSINK. — Distributional ecology of some Acrididae (Orthoptera) in the
dunes of Voorne, Netherlands, pp. 357—443, fig. 1, plates 2—5, 2 graphs.
| Tijdschrift voor Entomologie, deel 106, afl. 8. Gepubliceerd 2-XII-1963
| Nederlandsche Entomologische Vereeniging
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i
l
BRIVER
DISTRIBUTIONAL ECOLOGY OF SOME ACRIDIDAE
(ORTHOPTERA) IN THE DUNES OF
VOORNE, NETHERLANDS
BY
B. M. LENSINK
Oostvoorne
CONTENTS
General introduction
The vegetation of the landward dunes and the microclimate .
The vegetation .
Introduction
General description af the iaia
Botanical description of the vegetation .
Vegetational types and structural changes
The choice and distribution of the quadrats .
The microclimate .
The grasshoppers
The grasshoppers in the dec af ocre ped ‘the dute gi the os un
The identification of the instars .
The number of instars
The sampling technique 3
Periodicity and phenological data .
The numbers of grasshoppers .
The hatching sites .
Introduction i
The hatching sites of M. Ci È
The hatching sites of C. biguttulus
The hatching sites of C. albomarginatus ;
The distribution of the grasshoppers during the season .
The distribution of M. maculatus .
The distribution of C. biguttulus .
The distribution of C. albomarginatus -
The activities of the grasshoppers during the day .
Introduction u
The diurnal es of M. es
The diurnal activities of C. biguttulus
The diurnal activities of C. albomarginatus
Summary
Discussion .
Acknowledgments
References
358
360
360
360
361
362
363
370
371
374
374
376
DIET
378
379
384
387
387
387
392
396
398
398
411
417
427
427
428
434
435
437
440
442
442
358 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
GENERAL INTRODUCTION
Experience in making an inventory of a given region has consistently shown that
a species is not equally distributed over the area in which it occurs. In some places
it is completely absent and in others its numbers vary greatly from place to place.
In approaching this phenomenon from the point of view of the ecology of
distribution, attention is given on the one hand to the structure of the animal,
its physiological characteristics and behaviour, and on the other hand to the
environmental factors, both biotic and abiotic. At this point the student encounters
an extremely complex network of relationships. For the present it is impossible
to investigate the whole of such a network or even the whole of one of its parts.
We are forced to limit ourselves to the investigation of some of the relations
between some of these factors. This may, however, make it possible to distinguish
the nature of the totality of the mutually operative relations.
A systematic approach is nevertheless indispensable. This type of investigation
can usually be divided into three phases. First, the places in which a particular
species occurs in the area are established; second, in a field study the factors are
determined which might be responsible for the restriction of the species to that
environment; and third, a laboratory investigation is made to determine whether
the reaction of the animals to various factors is such that what has been observed
in the field can be understood.
The distribution of three species of grasshoppers, Myrmeleotettix maculatus
(Thunb.), Chorthippus biguttulus (L.) and Chorthippus albomarginatus (de
Geer), found on the island of Voorne (4°.5’ W, 51°.54 N) in the Netherlands
was investigated between 1953 and 1955. Of the three investigational phases
mentioned above, only two could be fully carried out. The newly-built field
laboratory was not yet properly equipped, which meant that the third phase could
only be realised for a few factors with very simple procedures. Nevertheless, this
situation did not detract from the fact that a comparative investigation into the
distribution of the three species of grasshoppers over a small area of the dune
region made it possible to understand the situation observed in the field, this in
turn providing an understanding of the nature of the relationships between the
various entities.
A study of the factors in the environment which determine localization requires
the analysis of the very complex totality of relationships affecting life. Such analysis
can only be productive when it is possible to select an environment without too
large variations or too strong extremes.
On the basis of these considerations, distribution was studied in a small area,
in this case a section of older stabilized (landward) dunes (measuring 1.5 hectare)
which was almost entirely covered with grass. This had advantages over a study
made in an area with a great variety of vegetational structures. It not only sim-
plified the technical aspects of the study but also permitted a more intensive study
of the mutual dependence of the distribution of the species in all their develop-
mental stages and the various environmental factors. In addition, the apparently
limited variation in this vegetation proved to be a crucial factor for organisms
spending their entire lives in that vegetation without many possibilities to escape
B. M. LENSINK : Distributional ecology of Acrididae 359
from it (UVAROV, 1928 and CLARK, 1948). This region had the further advantage
that the three species of grasshoppers are all present on the same terrain and could
thus be investigated simultaneously. Differences and agreement between phen-
omena offer the observer increased opportunity to penetrate to the basic relation-
ships between animal and environment.
This region was chosen after a rough inventory had been made of the dune
area on Voorne (p. 374).
A special sampling technique was developed for the investigation into the
distribution of the three species of grasshoppers, with particular view to the local
circumstances (p. 378). With this technique, the distribution could be studied
from the initial egg stage to the adult.
For this investigation a large number of quadrats was chosen in the various
types of vegetation and sampled during the season (from April to November)
with our method. Although no difficulty was encountered in identifying the adults
of the three species, practicable field-characteristics had to be chosen for distin-
guishing the immature stages of each of the species and the corresponding stages
of all three (p. 376).
During the three years in which the study was carried out, an attempt was made
to establish as accurately as possible when the various stages appeared in the field
and when they disappeared from it. The relationship between this annually recur-
ring phenomenon and the climatic factors at the time was also studied (p. 398).
In addition to the attention given to distribution during the season, observations
were also made of the activities of the animals during the period of a day under
various weather conditions (p. 427). It is particularly this latter kind of study
which, by observation of the animal from minute to minute, greatly increases the
understanding of an animal. It reveals the manner in which the animals move
about in the vegetation, where their courtship occurs, where and for how long
they sun themselves, what diet they choose, how and where the eggs are laid, how
they react to all kinds of enemies, etc.
In this picture, the vegetation is extremely important and very complex. It serves
not only as a source of food but also as a place in which to live. The vegetation,
which has a patchy distribution is characterized not only phyto-cenologically but
also structurally (p. 364). Special attention has been given to the structural changes
which occur during the year and from year to year. Periodic changes in the
vegetation produce changes in the character of the animal’s habitat. Such changes
may create conditions under which the animals cannot live, and this may result
in migration.
The vegetation is subject to climatic influences. But it is the vegetation and the
structure of the soil (which in its turn is also partially influenced by the vegeta-
tion) that create the peculiar phenomenon of the microclimate. VESTAL (1913)
refers to this point, and Uvarov (1928), FRANZ (1931), STROHECKER (1937)
and CLARK (1948), among others, emphasize the importance of microclimatological
investigation in the study of the distribution of grasshoppers.
The investigation of this extremely important microclimate could not be carried
out because the proper instruments were lacking. STOUTJESDIJK (1961) later
360 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
investigated, in the same locality, on a number of points the variations in micro-
climate within the vegetation in the course of a day and the season. These data
and the general pattern of the microclimate in the various types of vegetation are
described (p. 372). The investigation of the activities of the insects during the
day revealed very clearly how sensitive they are to changes in climatological
conditions. FRANZ (1931, 1933) stresses this sensitivity by calling the Orthoptera
climate indicators.
THE VEGETATION OF THE LANDWARD DUNES AND THE
MICROCLIMATE
THE VEGETATION
Introduction
The vegetation occupies an important place in the lives of the grasshoppers.
It is not only a source of food, it also forms a predominant part of the environ-
ment of these insects. For this reason, much attention was paid to the characteristics
of the vegetation present in the area in which the investigation was carried out.
As a source of food, the composition of the vegetation has little influence on
the distribution of the grasshoppers because their primary preference is for grasses
and sedges. These plants are present in the entire field in which the study was
made, some even dominating the grass cover.
As environment, however, the vegetation is very important for the distribution
of the grasshoppers. They seek those places that agree with their vital requirements
such as locomotion, protection against predators, protection against climatic
extremes, and the maintenance of reasonable heat and water economy.
This environment is determined by the structure of the vegetation and the
microclimate prevailing in it. These two aspects are very closely related. The
structure of the vegetation is determined by the qualitative and quantitative
characteristics of the various parts of which it is composed, and shows differences
from one place to another that are dependent of the botanical composition and
habitat factors. The structure of any given place will also in the majority of cases
vary rather strongly in the course of a year (seasonal dynamics, BOERBOOM, 1957a)
and may differ from year to year depending on annual changes in the macro-
climate.
During the investigation, as many data as possible concerning the vegetation
were collected and the most striking changes noted. Samples were made at many
points, and the development of the various layers in the vegetation and the height
of the most important plants was followed.
In the plant growth of the landward dunes, eight communities and vegetations
are distinguished. However, this characterization does not express the seasonal
dynamics of the vegetation. The vegetation, in all its variations, is therefore
B. M. LENSINK : Distributional ecology of Acrididae 361
described in a different manner which reveals peculiarities in the structure that
may be important to the life cycle of the grasshopper. On the basis of the finer
nuances in the vegetation, a number of types are distinguished, each with a specific
structure and annual development. For each of these types the grasshopper popu-
lation was estimated several times during the season in a number of 3 sq.m
quadrats.
The nomenclature used in this work for the higher plants is taken from HEu-
KELS & VAN OOSTSTROOM (1956) and for the mosses from VAN DER WIJK &
MARGADANT (1947). The infraspecific taxa of many plants characteristic for the
dunes were chosen, after consultation with E. VAN DER MAAREL who has investi-
gated the vegetation of the dunes of Voorne, in agreement with BOERBOOM
(1957b). The phyto-cenological nomenclature is borrowed primarily from WEST-
HOFF, DIJK, PASSCHIER & SISSINGH (1946).
General description of the terrain
The grassy and slightly rolling landward dunes on the border of the munici-
palities of Oostvoorne and Rockanje comprise two parts separated by market
gardens, hedges and a road. The investigation was carried out in the part of the
terrain lying furthest to the west (Plate 2). This part of the dunes is bordered on
the west, north, and east by market gardens. The only connection between this
part and the younger dunes is by a small path through a wood. The dunes are
separated from the market gardens by a thick and in some places quite high hedge
of hawthorn trees. To the south is the Tenella-plas, an educational public garden
surrounded by trees of medium height. The entire terrain is therefore completely
isolated from all other dune regions in which grasshoppers are found. Migration
from these regions is practically excluded; grasshoppers are seldom seen in the
market gardens.
The maximal difference in elevation between the highest and lowest ground is
3 metres. The more elevated parts of the otherwise rather flat field are relatively
dry. The lower parts are rich in humus and moist; particularly in the winter and
early spring of the years in which the investigation was done, these places were
spongy and in some spots even covered with water.
The landward dune formerly served as pasture land for goats and horses, and
this fact probably gave the present vegetation the chance to develop. A large
number of plant species is found, more than 60 species being present.
The area may be generally characterized as dry and more or less densely covered
grassland. Wide differences in structure occur, partially caused by the stand. The
higher parts are covered with a thin vegetation, often rich in herbs, with a highly-
developed moss layer that partially or entirely covers the soil. Where the moss
does not entirely cover it, the sandy soil is visible. In the transitional areas between
the higher and lower parts, the grass cover is thicker and the moss covers less of
the surface. In the lower spots the grass cover is very thick and locally rich in
herbs. In late spring the grass in this vegetation becomes fully developed.
Locally, Salix repens penetrates the grass cover from the hedges. A high, thick
growth of grass is sometimes found between the low shrubs.
362 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Botanical description of the vegetation
The vegetation of the terrain in which the investigation was carried out
comprises a complex of communities belonging primarily to the Koelerion albes-
centis, but also in part to the Corynephorion canescentis.
The characteristic species and the species occurring frequently among the Koele-
rion are: Lotus corniculatus var. ciliatus, Taraxacum rubicundum, Ranunculus
bulbosus, Ononis repens var. mitis, Erigeron acer, Arenaria serpyllifolia, Erodium
glutinosum, Phleum arenarium, Thymus pulegioides, Plantago lanceolata vat.
Sphaerostachya.
In addition there are few species from the Bromium erecti: Potentilla taber-
naemontani and Helictotrichon pubescens.
The species from the Corynephorion are: Corynephorus canescens, Aira praecox,
Polytrichum juniperum, Festuca ovina, Polygala vulgaris, and Poa pratensis var.
humilis.
A few different species are: Galium verum var. maritimum, Carex arenaria,
and Leontodon nudicaulis.
In this complex, eight communities and vegetations are to be distinguished.
They are to a great extent determined by the stand.
(1) Tortuleto-Phleetum arenarii, an open vegetation on the south slope with
an incomplete moss layer of Tortula ruraliformis and Phleum arenarium, Erodium
glutinosum, E. cicutarium, Myosotis hispida, and Veronica arvensis. Two transitions
of this community are found:
(a) A transition to Taraxaco-Galietum, consisting of a vegetation with a half
closed to entirely closed moss layer of Hypnum cupressiforme, in which
Galium verum predominates, and
(b) A transition to Violeto-Corynephoretum dunense, in which tussocks of Co-
rynephorus canescens occut.
(2) Taraxaco-Galietum, a vegetation with a closed moss layer of Hypnum
cupressiforme with Galium verum var. maritimum, Thymus pulegioides, Poa pra-
tensis var. humilis, Lotus corniculatus var. ciliatus, Carex arenaria, and Festuca
rubra subvar. arenaria. The herbs form a low, dense and closed layer. This
vegetation occurs on the slightly higher parts of the terrain.
(3) Festuceto-Galietum maritimi. Of these a poor form occurs with Festuca
ovina in tussocks and rosette plants of Leontodon nudicaulis, Hieracium pilosella,
and a moss layer of Hypnum cupressiforme and Cladonia species. This vegetation
covers a large flat part of the southernmost part of the terrain.
(4) An incompletely developed Violeto-Corynephoretum dunense, without
Viola canina and with few Cladonia species. This vegetation is found in only a
few places.
(5) A vegetation of Sedum acre and Plantago coronopus. This is found along
paths and around depressions. It probably develops where calcareous sand has
worked its way to the surface.
(6) A vegetation of Festuca rubra subvar. arenaria and Carex arenaria, a form
with Calamagrostis epigejos; in this vegetation Calamagrostis can dominate so
B. M. LENSINK : Distributional ecology of Acrididae 363
strongly locally that facies-formation occurs. This vegetation occurs only locally
and spreads from the borders of the terrain.
(7) A vegetation of Sieglingia decumbens and Gentiana campestris baltica.
This form of vegetation is found at the transition from the higher parts to the
lower. Grasses such as Sieglingia decumbens, Anthoxanthum odoratum, and Holcus
lanatus predominate here, forming a grassy cover on rather moist ground.
(8) A vegetation with Lythrum salicaria and Lysimachia vulgaris. This vege-
tation grows on the moist soil, rich in humus, of the lowest parts of the terrain.
It often forms a growth rich in herbs with such species as Achillea millefolium,
Potentilla erecta, Hydrocotyle vulgaris, Potentilla anserina, Holcus lanatus, Sieg-
lingia decumbens, Anthoxanthum odoratum, Plantago lanceolatum var. sphae-
rostachya, and Carex flacca.
Vegetational types and structural changes
A list of the plant-communities and vegetations found in the landward dunes
will give plant ecologists and biocenologists a clear impression of the nature of
the overgrowth, but as a description of the habitat of the grasshopper it is not
adequate. Even the samples of the various vegetations would fail to provide a real
insight into the habitat and the changes taking place in it. A more detailed de-
scription of the structure is required. This point is often neglected by animal
ecologists. Several botanists have offered a type of structural description which has
very attractive aspects for the field ecologist. DANSEREAU (1951) made use of
symbols expressing many characteristic peculiarities of the plants in a vegetation.
The structure of the vegetation is thus symbolized by a diagram which, however,
fails to reflect the natural situation. Each symbol reflects the relative space occupied
by a plant type in the community. It serves both for individuals of one species
and for individuals of various species. Such a diagram, however, always requires
an accompanying description.
Although the value of this method was not underestimated for the present
study (it would, for instance, be extremely serviceable for comparisons of vege-
tations at different geographical latitudes), it was considered to offer too few
possibilities in a physiognomically simple vegetation such as the growth on the
landward dunes. The use of symbols as proposed by DANSEREAU provides an
over-generalized picture and also fails to reflect periodicity.
For the present study, in characterizing the vegetation of the field a detailed
description is first given of the various types of growth to be distinguished in it.
This is illustrated by a number of drawings of these types (fig. 1). These drawings,
however, refer exclusively to the natural situation in the month of June.
A structural description requires in the first instance the distinguishing of as
many vegetational layers as possible and the indication of the cover in per cent.
In consultation with E. VAN DER MAAREL who made a study of the structural
description from the botanical point of view, the following surface vegetation
layers were tentatively distinguished: the moss layer, the rosette layer, the carpet-
ting layer, the high herb layer and the low shrub layer.
364 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
These layers may be briefly described first as follows.
(1) The moss layer. The mosses often cover a large part of the soil; in denser
vegetations they remain more in the background.
(2) The rosette layer. Under this term is understood the layer of herbs that
form rosettes in the moss layer and barely project above it, such as Hieracium
pilosella, Taraxacum spec., Hypochaeris radicata, Leontodon nudicaulis and typical
low-growing creepers such as: Potentilla tabernaemontani and Ranunculus repens.
Only the flower stalks of these species project above this layer.
(3) The carpetting layer. A large number of herbs show a strong horizontal
spread and form a low cover above a highly developed moss layer. These species
are: Lotus corniculatus var. ciliatus, Galium verum var. maritimum, Polygala vul-
garis, Ononis repens var. mitis, Cerastium holosteoides, Cerastium arvense, Vero-
nica arvensis, and a few creeping species such as: Thymus pulegioides and Sedum
acre. Between these herbs a few of the low grasses such as Agrostis stolontfera
and Bromus mollis var. nanus occur.
(4) The high herb layer. This layer is formed by the more vertical plants, the
grasses in particular dominating. The layer includes: Festuca rubra subvar. arenaria,
Festuca ovina, Corynephorus canescens, Sieglingia decumbens, Calamagrostis
epigejos, Carex arenaria, Helictotrichon pubescens, Poa pratensis var. humilis,
Anthoxanthum odoratum, Luzula campestris, Holcus lanatus, Carex flacca, Achillea
millefolium, and Erigeron acer. This layer occurs in various forms. A number of
grasses such as Festuca ovina and Corynephorus canescens grow in tussocks, which
creates a characteristic structure. Other grasses and Carex arenaria influence the
structure of the vegetation by their dominancy. In addition, differences in the
density of this layer are caused by variations in the amount of horizontal spread
of the species present.
(5) The low shrub layer. This layer is formed by Salzx repens, small shrubs
40 to 80 cm in height. Locally, and in the summer barely projecting above the
high herb layer, Rosa pimpinellifolia (25—40 cm) occurs.
On the basis of typical differences in the share taken by these layers in the
vegetation, I have distinguished seven structural types. Type I through type VI
are grass vegetations, type VII comprises the Salix repens shrub that spreads into
the terrain from its borders and between which a dense grass cover is sometimes
present. Type VII was not sampled systematically.
Type I. The vegetations belonging to this structural type are open and low
(Plate 3). The moss layer covers the soil only partially (coverage 0—10%);
Hieracium pilosella very locally, sometimes forms facies. The herbs appear only
to a small extent, and the sparsely distributed grasses and herbs in the high herb
layer cover the soil to a very limited degree. Festuca rubra subvar. arenaria and
Carex arenaria dominate in this layer. Locally, tussocks of Corynephorus canescens
occur on an otherwise very scantily covered soil.
In the course of the year, the aspect of the vegetation changes appreciably. The
period of the greatest vegetative expansion falls in the winter and the early spring.
The moss cover is then saturated with moisture and as a result shows a fresh green
colour. The superficial sandy soil is moist and firm. A number of annual herbs
B. M. LENSINK : Distributional ecology of Acrididae 365
occur in the moss layer (winter annuals), such as Erophila verna, Saxifraga tridac-
tylitis, Aira praecox, Phleum arenarium, and Vicia lathyrotdes. The perennial
species are still small and of a yellowish-green colour. The green grass cover is
S— 10 cm high. Scattered over the soil are dead leaves of grass and flower stalks.
Later in the spring, in April and May, the winter annuals flower and fructify.
They are not numerous and most of them barely project above the moss layer.
In this period the remaining herbs develop.
In the early summer the bare sandy soil becomes dry and develops a crust, or
where it is walked over, is very loose. The moss cover also becomes very dry,
changes to a yellowish-green colour and shows splitting. Of the winter annuals
only dried remnants can be found. Some of the other species come to the fore.
Rosette plants develop and creeping plants spread horizontally over the moss cover.
In the summer during flowering, some of these plants add colour to the vegetation.
The grasses and the sand sedges flower and fructify with stalks as high as about
20 cm. Their vegetative part does not grow higher than 10 cm. A few plants such
as Senecio jacobaea shoot up very high during the flowering period.
In late summer and in the autumn, more and more dead leaves and stalks appear
and are pressed down on the moss cover by the wind. The mosses become greener
again and the winter aspect gradually asserts itself. The vegetations described here
occur on the southern slopes and belong to transitions from the Tortuleto-Phleetum
arenarii to the Taraxaco-Galietum maritimi and the Corynephoretum dunense.
They also occur, distributed over the field, in places where the vegetation was
damaged before and during the Second World War, having originally belonged
to the Taraxaco-Galietum maritimi.
Type II (Plate 3). The vegetations of this type have a well-developed moss
cover (ceverage: 60—100%). The soil is entirely covered by the mosses and herbs
so that the sandy surface is not visible. In the rosette layer many rosettes of Hzera-
cium pilosella are found, a species which is often extensively represented in this
type and even forms facies, in which case its coverage is 50—100%. The carpet-
ting layer is better developed than in type I, especially Galium verum var. mari-
timum and Thymus pulegioides predominating. The grasses Festuca rubra subvar.
arenaria, Agrostis tenuis, and the sedge Carex arenaria form a thin layer to a
height of about 10 cm.
The appearance of type II changes less in the course of the year than that of
type I. The moss layer is saturated with moisture in the winter and early spring.
Winter annuals are far less frequent. In the early summer the mosses become dry
but continue to form an unbroken layer. Locally, the activity of ants brings some
sand to the surface. The remaining species in the carpetting layer develop and
flower. They cover only a small part of the surface, however (— 50%). The
high herb layer becomes higher during flowering because of the flower stalks
(20 cm). In the autumn many dead leaves and stalks are seen on the moss layer.
These vegetations occur on the highest parts of the terrain and may be assigned
to transitions from the Tortuleto-Phleetum maritimi to the Taraxaco-Galietum
maritimi.
Type III. The vegetations belonging to type III correspond closely to the over-
growth described for type II. Instead of free-standing grasses in the high herb
366 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
layer, tussock grasses are predominantly seen, which creates the typical structure
of this type (Plate 4).
The moss layer covers 30—50% of the surface; the rosette layer and the carpet-
ting layer do not grow higher than 5 cm and cover 30—60% of the surface.
In the high herb layer tussocks of Festuca ovina occur, covering 30—50% of the
surface. In the period in which the grass flowers, a great number of flower stalks
grow up above the vegetative part of the plants (10 cm), to a maximum height
of 25 cm.
These plants cover a large part of the terrain and belong to the Festuceto-
Galietum maritimi.
Type IV. In the moderately dense to dense low vegetation, the moss layer is
well developed (coverage: 60—100%). A rosette layer is almost entirely absent:
it blends into the carpetting layer which is dense but not higher than 5 cm. Both
layers combined cover 60—100% of the surface. They include: Lotus corniculatus
var. ciliatus, Galium verum vat. maritimum, Thymus pulegioides, Cerastium cf.
tetrandum, Plantago lanceolata vat. sphaerostachya, Potentilla tabernaemontani,
Luzula campestris, and Achillea millefolium. The high herb layer constitutes a
grass cover covering a maximum of 30% of the soil. The vegetative parts are
10—15 cm high, the flower stalks growing up 20—30 cm. A few of the species
in this group are Festuca rubra subvar. arenaria, Agrostis tenuis, Carex arenaria,
Festuca ovina, Poa pratensis var. humilis, and Achillea millefolium. In various
places along the borders of the terrain or near vegetations composed of a high,
thick grass cover, the flower stalks of Calamagrostis epigejos rise up to a height
of 40 cm in this type.
The changes in the vegetation over the year are again less marked than for the
previously described types. In the course of the summer the cover provided by a
number of the most frequently occurring species in the carpetting layer is in-
creased; the plants in the high herb layer begin to flower. The latter cause little
change in the structure of the very dense carpetting layer.
The vegetation of this type belongs to the Taraxaco-Galietum maritimi and
occurs over a large surface on the higher parts of the terrain.
Type V. The vegetation included in this type are distinguished by the higher
proportion of grasses in the high herb layer. The structure of the higher layers is
consequently much denser than in the corresponding layers of the other types.
The moss layer is of little importance. Rbytidiadelphus squarosus predominates.
There is no rosette layer, the rosette plants do not spread their leaves horizontally
on the soil. The carpetting layer shows a transition to the high herb layer.
Two different forms can be distinguished within this type. The first is a
vegetation occurring on the transitional slopes between the highest to the lowest
parts of the terrain. It consists of a moderately dense to very dense, gentian-rich
grass cover, in which among others are present: Anthoxanthum odoratum, Sieg-
lingia decumbens, Festuca rubra subvar. arenaria, Holcus lanatus, Agrostis tenuis,
and Gentiana campestris baltica.
The vegetation covers 80—100% of the surface and its vegetative part is
10—15 cm high; in the flowering period the many flowering stalks may reach a
height of up to 25 cm. Locally, a vegetation is also found that is rich in Rosa
B. M. LENSINK : Distributional ecology of Acrididae 367
pimpinellifolia, small shrubs 25—40 cm high; these roses, which bear numerous
small leaves in summer, cover 80% of the surface and cast much shade on the
underlying layers. The higher herb layer covers 20—100% of the soil. Many
plants occur in this layer: Achillea millefolium, Cerastium tetrandum, Luzula
campestris, and a number of rosette plants such as Ranunculus bulbosus, Taraxa-
cum spec., and Plantago lanceolata var. sphaerostachya. The grasses Helictotrichon
pubescens, Festuca rubra subvar. arenarta, Agrostis tenuis, Calamagrostis epigejos,
and Carex arenaria are numerous between the dune roses.
In winter and spring this vegetation, excluding the dune roses, is 15—20 cm
in height and consists of the vegetative parts of the herbs. In spring the grasses
grow quickly, especially Helictotrichon pubescens. The weather conditions pre-
vailing in this period are of great importance for the extent of this growth. Most
of the herbs do not grow much higher than 20 cm, with the exception of a few
flower stalks (Achillea millefolium). In the summer and autumn the dry stalks
of many grasses, which have already finished flowering, fall to form a dead layer
in the high herb layer. The dune roses shrubs are leafless in the winter, in the
summer they come more to the fore in and above the high herb layer. It is striking,
lastly, that in some parts of this vegetation the structure is less dense because of
the large numbers of rosette plants that spread their leaves between the grasses.
Type VI. Type VI includes the vegetation of the lowest parts of the terrain
and its edges along the hedgerows. It comprises densely covering grass vegetation,
rich in herbs in the parts with the most moisture (Plate 5).
The moss layer is either absent or very fragmentary. The soil is covered with a
layer of dead grass and flower stalks, sometimes as much as 10—15 cm thick.
The carpetting layer on the wettest spots is 10 cm high and covers 60—100%
of the surface. It includes: Achillea millefolium, Ranunculus bulbosus, Taraxacum
spec., Prunella vulgaris, Plantago lanceolata var. sphaerostachya, Galium uligino-
sum, Potentilla anserina, Lotus corniculatus, Hydrocotyle valgaris.
Growing to a height of about 40 cm in the high herb layer are: Lythrum salt-
caria, Lysimachia vulgaris, Pulicaria dysenterica, and Achillea millefoltum. The
grasses that frequently occur in this layer are: Holcus lanatus, Calamagrostis
epigejos, Festuca rubra subvar. arenaria, Agrostis stolonifera, Steglingia decumbens,
Anthoxanthum odoratum, and the sedges Carex flacca and Carex arenaria.
On higher spots in the terrain a high, dense grass cover is found locally in
which Calamagrostis epigejos dominates. The non-grassy herbs are forced into the
background. This vegetation shows strong growth in the summer and its flower
stalks reach a height of a metre in some cases. The soil is covered with a 20 cm
thick layer of dead plant material, only the surface of which dries out in summer.
The appearance of this vegetation varies greatly, due especially to the grasses.
At first the vegetative parts of the plants are not higher than 20 cm, but in the
course of the summer some species show rapid growth to an appreciable height
as a result of which the structure becomes very dense. In the autumn and winter
most of the grasses collapse and their dead parts accumulate on the ground.
Type VII. The dwarf shrubs, consisting of Salix repens and a moderately dense
to dense grass cover, comprises this type of vegetation. The shrubs reach a height
of 40—80 cm with a coverage of 60—100%. The grasses grow rapidly in early
368 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
type IV
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369
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370 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
summer, causing a strong increase in the density of the vegetation among the
shrubs.
In addition to the developmental changes in the various layers of the vegetation
in the course of the year, differences are observed from year to year in the ap-
pearance of the vegetation. For instance, in May and June, 1955 the grasses showed
a more luxuriant growth than in the same months in 1953, 1954, and 1956. This
vigorous growth was probably induced by heavy rains, in particular in May, 1955.
Although the grasses had developed more fully throughout the terrain in which
the investigation was carried out, the structural differences were strongest in the
vegetations belonging to types V and VI, which occur in the driest areas. As a
consequence, in 1955 the structure of these vegetations was high and dense at an
earlier date than in other years, and in addition the grass cover spread towards the
less dense vegetation. It is self-evident that observations of such vegetational
changes from year to year ate very important to the estimation of the distribution
of the grasshoppers.
Changes in the vegetation were also established in succeeding years. In addition
to a marked spreading of the shrub, Calamagrostis epigejos was observed to show
local predomination.
The increase in shrub components on the landward dunes accompanies a loss of
characteristic peculiarities of the landscape.
The data concerning the description of the vegetation are arranged in Table 1,
in order to give a general impression. An illustration of the various types is given
in fig. 1, the drawings representing the situation in the month of June.
The choice and distribution of the quadrats
The choice of the quadrats in the early summer of 1953 was based on the dif-
ferences in the structure of the vegetation. Fifty quadrats measuring three sq.m
were selected, and the grasshopper populations were investigated in 1953, 1954,
and 1955. In 1955, another group of quadrats was laid out for more detailed
investigation of certain activities of the grasshoppers. For each of the types of
vegetation, the quadrats were chosen in the most densely populated locations.
In 1954, 55 additional quadrats measuring one sq.m were sampled for the in-
vestigation of the hatching sites. Table 2 gives the number of three sq.m quadrats
systematically sampled from 1953 through 1955 in each type of vegetation.
Table 2. Number of quadrats sampled from 1953—1955.
Type I ee V | VI |
1953—1954
1955
quadrats
quadrats
Two of the group of 50 quadrats were found to be transitions between two
types and two other quadrats were later found to lie in an area very frequently
visited by people. These four quadrats were excluded from the investigation.
B. M. LENSINK : Distributional ecology of Acrididae 371
THE MICROCLIMATE
The microclimatological conditions are important not only for the rate of
development of the insects and their condition in the various stages, but also for
all the activities of the insects. In the places where grasshoppers occur, attention
must be given to such factors as temperature and moistness of the immediate
surroundings in addition to the factors of the macroclimate. Lack of suitable in-
struments and the impossibility of obtaining them in time for the investigation,
and the fact that the taking of the samples proved to require all the available
working time, prevented serious investigation of the microclimate. However,
starting in 1956, at Weevers’ Duin Biological Station in Oostvoorne, Dr. PH.
STOUTJESDIJK carried out microclimatological measurements for biological pur-
poses in many types of vegetation found in the dunes of Voorne (STOUTJESDIJK,
1961), including the terrain used in the present study. Using these data and others
from the microclimatological literature (BRUNT, 1945, GEIGER, 1950, SUTTON,
1953), a general discussion can be given concerning the course of the microclimate
in the various types of vegetation, as distinguished by me, in the periods of the
year that are of importance for the grasshoppers. An attempt is also made to
roughly characterize the ecological factors of the environment.
In general it may be stated that the various microclimatological factors manifest
themselves quite differently above a soil surface covered with vegetation than
above a surface with no plant cover. Even a very sparse growth introduces changes
that moderate the climate immediately above the soil. In this sense, for example,
a surface with no plant cover may reach very high temperatures on warm, sunny
days, while the same values are appreciably lower when some cover is present.
This factor alone is sufficient to produce wide direct and indirect differences
in the microclimate.
The microclimate is formed by the influence of the climate as a whole, by the
nature of the soil, exposure, and all kinds of particularities of the vegetation.
The combinations of species and the height and density of the plants are of im-
portance. The denser the vegetation, the cooler it is during the day near the ground.
The solar radiation is taken up by the leaves and stalks of the grasses and other
plants, creating deep shade in the layers below. The thick absorption layer (soil
and vegetation) distributes the heat in such a way that high temperatures do not
occur as they do immediately above a bare or sparsely covered soil. A thick grass
cover with a layer of dead leaves contributes to the occurrence of low nocturnal
temperatures within and above it, especially on clear, windless nights with high
radiation. This cooling process is accentuated by the slightness of the horizontal
air currents and the fact that little heat is radiated by the soil.
In the terrain in which the study was carried out the microclimate is subject to
great variation caused by differences in elevation as well as by the vegetation,
which shows distinct structural differences from place to place. The grasshoppers
which live in this vegetation must therefore make a choice during the season and
in the course of a single day throughout their development. The following material
contains some considerations concerning the microclimate in the various types of
vegetation distinguished in this study.
372 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Type I. The vegetations included in type I are very thin in structure; above an
incompletely developed moss cover there is a very scanty grass cover. The micro-
climate in this extremely open vegetation often shows extreme variations. On
sunny days, a large part of the solar radiation is absorbed by the surface of the
soil. Under favourable conditions high surface temperatures can develop, which
lie 20—30° C above the air temperature at a height of two metres (STOUTJESDIJK,
1959, 1961). In addition, as a consequence of the limited flow of moisture from
this type of vegetation, the relative humidity of the air is low.
One cm above the ground the temperature is appreciably lower. STOUTJESDIJK
states that the air temperature one cm above the soil is even much lower than the
soil temperature at a depth of two cm. It was also found that directly above the
surface there is very wide fluctuation in temperature. In addition, after sunset the
temperatures of the surface layers of the soil drop sharply; the sand can become
very cold, especially on very clear, windless nights. It is in just this layer with
wide temperature variations that the grasshoppers occur.
In general, the nocturnal temperature of the air layers immediately above the
soil and of the soil itself remains a few degrees higher than the temperature at a
height of two metres, which points to heat radiation by the soil. The same picture
holds in general for the spring season, although the absolute values lie lower than
in the middle of the summer.
Type II. The vegetations of type II differ from type I in the development of
the moss layer. The entire soil surface is covered with a well-developed moss cover.
The microclimate in this type shows strong agreement with that in type I, but in
this case the moss layer, which dries rapidly, absorbs most of the radiation and
therefore reaches high temperatures. The soil temperature under the moss is ap-
preciably lower.
Type III. The vegetations of type III are characterized by tussocks of grasses
between which a more or less closed moss cover occurs. The microclimate above
the moss cover is comparable to that of type II, although the values may lie some-
what higher because the cooling effect of the wind is greatly reduced by the grass
tussocks. In the tussocks themselves the highest temperatures are found just under
the tops of the plants, because the solar radiation is taken up by the leaves and
flower stalks of the grasses. The temperature will be only a few degrees higher
than the air temperature just above the plants. The tussocks also cast shade on the
moss cover, especially when the sun is lower in the sky. In this type of vegetation,
consequently, opposite extremes may occur quite close together, i.e., maximum
temperatures may occur as high as those occurring on the surface in Re II, but
the average temperatures lie lower.
Type IV. The vegetations of this type are open (star rich in grass and other
plants, in which rosette plants occur frequently in the approximately five cm thick
carpetting layer. Due especially to the presence of the rosette plants, the solar
radiation can penetrate deep into the vegetation locally. Since the grasses and herbs
decrease the drying effect of the wind above the moss cover, high temperatures
occur in some spots. The local humidity of the air is high as a result of the high
transpiration of the plant cover and evaporation from the soil, which has a
relatively high humus content. Shady spots occur beside many plants, with the
B. M. LENSINK : Distributional ecology of Acrididae 373
result that in this vegetation too, mild conditions are found next to heat.
Type V. The vegetations of type V are rich in grasses and herbs, with low,
closed growth. Solar radiation penetrates the vegetation less deeply. The highest
temperatures, which are not very high because of the strong transpiration of the
many leaves, are found above the soil between the tips of the leaves. Deeper in
the vegetation the temperature is lower, and a relatively large amount of shade is
created by the layer of leaves above. Later in the year there is more shade because
the sun is lower in the sky; the temperatures in the vegetation are lower than in
July, even during the warmest part of the day.
In the early morning, dew often forms in this vegetation, because the moist air
between the higher grass exposed to the cooling effect of the radiation does not
receive sufficient heat from the lower layers. This phenomenon is not seen in the
more open growths of types I, II, and III until later in the season.
Type VI. Type VI includes the high, dense grass vegetation found especially in
the lower areas of the terrain. A number of remarkable observations have been
made by STOUTJESDIJK (1961) in this type of vegetation on the same terrain. In
a closed, high stand of Calamagrostis epigejos varying between 45 and 75 cm in
height, the highest temperatures were recorded at the level of the lowest of the
leaf tips. Most of the solar radiation is absorbed there and converted into heat and
evaporation energy. On 10 July, 1959, with an air temperature of 22°C a tem-
perature of 24° C was measured between the tops of the leaves; the temperatures
in this vegetation are therefore not very high. The soil is much cooler and receives
heat from the top layer. On the afternoon of the same day (reduction in the rad-
iation intensity and more shade in the vegetation), the temperature gradient was
still slight and the fluctuations in temperature were also small. During the clear
evening which followed, however, the lowest temperature was found at the level
of the lowest tops (reading at 9:10 p.m.). Both above and below this level the
temperature was several degrees higher. The wind velocities in this dense vege-
tation are very low.
In less uniform vegetation of this type, consisting of tussocks of Calamagrostis
epigejos between which layers of flattened grass or low herbs occur, the tem-
perature profile is very different between the tussocks. The highest temperature is
measured a few millimetres above the surface of the dead leaves. On 11 July,
1959, at 12:00 noon the average local temperature was about 23° C higher than
at a height of one and a half metres. The fluctuations in temperature are very
large: STOUTJESDIJK observed temperature extremes at a height of three mm,
ranging over a 15° C difference during an observation period of about one minute.
The fluctuations remain large at even greater heights between the tussocks.
On still, clear nights the vegetation takes heat from the air, causing a drop in
temperature of about 30° C at a height of 3 mm, while at a height of one and
a half metres the temperature drops only a few degrees. The formation of dew
draws a great deal of moisture from the air, and this withdrawal of heat together
with moisture during the entire night causes very low temperatures and vapour
pressure to develop. On 12 June, 1959, at 4:07 a.m. a temperature of — 7° C
was measured at the soil surface as a result of very favourable conditions: the dry,
374 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
rather cold air on a clear, still night; the inability of the cold air to escape from
the low spots in the terrain; and, perhaps the most important factor, the unusual
heat economy of the soil which is covered with an insulating layer of dead grass.
This situation occurs in many of the spots where quadrats were chosen for
sampling. In May, low temperatures are still recorded in the vegetation, and it is
self-evident that such microclimatological conditions have very definite consequen-
ces for the grasshoppers.
THE GRASSHOPPERS
THE GRASSHOPPERS IN THE DUNES OF VOORNE AND THE CHOICE OF THE
WORKING TERRAIN
Investigation of the Acrididae in the dunes of Voorne showed that eight species
occur. These species are: Myrmeleotettix maculatus (Thunb.), Chorthippus bigut-
tulus (L.), C. brunneus (Thunb.), C. mollis (Charp.), C. albomarginatus (de
Geer), C. parallelus Zett., Omocestus viridulus (L.), and Oedipoda coerulescens
(L.).
An understanding of the distribution of these species indicates why a given area
with a given number of species was the obvious choice as working terrain for this
investigation. The density of the population and the variegation of its composition
are factors affecting this choice. The density and variegation depend to a large
extent on the vegetation. In this connection it has been found (VAN DER MAAREL,
1960) that the amount of shrubs in its various developmental phases, with its
heavy to medium heavy growth, excludes a grasshopper population in many places.
The small grassy areas occurring here and there between the shrubs are too small
to permit more than a limited grasshopper density. Open spots, thinly covered
dune slopes, and the verges of the many foot-paths are also lightly populated.
The most suitable environment is offered by terrains with a grass cover, such as
are found in the landward dunes.
The inventory made in 1953 supplied the following data.
(1) Myrmeleotettix maculatus occurs in varying densities throughout the dune
region. In the small open spaces between the Hippophae-shrubs and along the
paths, the numbers are small. On the coastal ridge, a few specimens are found
here and there; the local climate, being extreme, apparently prevents a population
increase. The landward dunes, with extensive grassy surfaces, show the highest
density. In the direction of the polder, M. maculatus is rarely found because of the
increasing amount of woodland and farms.
(2) Chorthippus biguttulus occurs only locally, especially in the old dunes, in
the low, thick grass cover of the driest areas and in low Salix repens and Hip-
pophae shrubs or near them.
(3) Chorthippus brunneus is distributed over the entire dune region, but in
limited numbers. It prefers dry, sunny places such as those along the paths in the
dunes. This species is frequently seen in the grass verges of the roads in the land-
ward dunes and in ruderal places. If only this grasshopper is present it is
B. M. LENSINK : Distributional ecology of Acrididae 375
found in only a few spots. Frequently, C. brunneus and C. biguttulus are present
simultaneously. PERDECK's experience (1957) concerning the choice of habitat
by these species agrees completely with mine. C. biguttulus prefers denser and
often somewhat more humid vegetations than C. brunneus.
(4) Chorthippus mollis is found in the dune region locally; this species lives
in the dry, dense vegetations in which a low Hippophae-shrub, Salix repens and
Ligustrum vulgare form an element. C. mollis is often found as sole species in the
dunes of Oostvoorne. Here and there this species lives, like C. brunneus, together
with C. biguttulus (but in limited numbers).
(5) Chorthippus albomarginatus lives in the dense, often high grass cover of
the dune meadows in the relatively damp parts of the dunes. It apparently prefers
the old parts of the dunes over the younger ones. It increases in number in the
direction of the polder, and is often seen in the polder itself.
(6) Chorthippus parallelus is found in large numbers in only one place, namely
in a meadow in the woods called the Mildenburg, in the municipality of Oost-
voorne. Outside the dunes this species is found on the ramparts of Brielle. Up to
the present it has not been observed between the two places.
(7) Omocestus viridulus was observed only once (in 1954) in an Agropyretum-
vegetation on the Kruininger Gors along the Brielse Maas river.
(8) Oedipoda coerulescens is seen in the dunes only in small numbers; it
prefers warm, dry places.
It is apparent that, in connection with the vegetational factors discussed in the
preceding chapter, the density of the grasshopper population in the dune region
proper is limited. The grassy landward dunes on the border between Oostvoorne
and Rockanje, however, show larger concentrations, particularly of M. maculatus,
C. biguttulus, and C. albomarginatus. These landward dunes were chosen for the
working terrain, a choice which offered many advantages. In the main, the three
species are the only ones occurring there, and the nymph stages of these species
(with the exception of the first instar) are easily distinguished from each other.
In addition, the vegetation of this terrain is extremely suitable for quantitative
sampling. Its location is also very favourable, particularly with respect to the
Weevers’ Duin Biological Station.
In very small numbers, three other species occur: C. brunneus, C. mollis, and
O. coerulescens. C. brunneus was observed on only a few occasions and during the
three-year study C. mollis was seen only once (1954), in a part of the landward
dunes lying outside the working terrain. O. coerulescens occurs only on thin
vegetation; it is easily distinguished so that confusion with the other species is
excluded. The instars of C. brunneus and C. biguttulus are difficult to distinguish
from each other in the field. Because the latter occurs on this field in such small
numbers, the doubtful cases were very limited in number and did not affect the
quantitative investigation.
In 1953 two grasshoppers were heard and seen whose song deviated from that
of C. biguttulus. PERDECK investigated one of these and identified it as a hybrid
(C. biguttulus x C. brunneus). The song contained elements of that of C. brun-
neus.
376 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
THE IDENTIFICATION OF THE INSTARS
At the beginning of the investigation, only a guess could be made as to which
species of grasshoppers inhabited the chosen terrain. In order to become familiar
with the immature stages of these species as rapidly as possible, nymphs were
collected in the field directly after hatching and raised in the laboratory to adult-
hood. Experience has shown that observation of the insects during rearing is
absolutely necessary for recognition in the field. Descriptions in literature can
convey only an impression. The appearance in 1954 of RICHARDS & WALOFF's
publication provided an opportunity to check my experience against their table
for the identification of the various instars and their determination table for the
nymphs of a large number of species, including M. maculatus and C. albomar-
ginatus. My experience is in agreement with the characteristics established by these
authors for both species.
However, for the recognition of the individual instars, their material gave little
support for my work. Measurements and counts of the insects in the field cannot
be made if it is necessary to release the nymphs directly after ward. The nymphs
in the field were not distinguished as to sex, since in the first instar this requires
too close an examination during which it is almost impossible to avoid damaging
the insects and consequently causing an unnecessary increase in mortality.
The rearing of nymphs is not easy. It often involves high mortality, which
requires beginning with large numbers. The collection of the first instar in the
field must consistently be done with the greatest care, especially because of their
fragility. Catching them with the hand or sweeping net damages them so badly
that only a few survive. They can best be caught in a glass tube. The tube is placed
over the nymph, which usually jumps up into the tube. The insect can then be
transferred to a glass bottle, the bottle being filled with dead grass to prevent the
insect from killing itself by jumping against the glass. Direct sunlight must be
avoided, because of the fatal effects of too much heat and moisture. When such
precautions are taken, laboratory culture is much more successful: the mortality
drops to about ten per cent.
For rearing the nymphs are transferred to a glass cylinder, ten cm long and
two cm in diameter, open at both ends. One end is covered with gauze and the
other plugged with a cork. Fresh grass must be fed daily. Before the last instar
moults, the insects must be transferred to a larger tube with a diameter of four
cm, containing a twig. Rearing experiments done by Miss A. S. C. SCHIJFSMA in
Leiden showed that the last mould is more successful if the insect can inflate its
wings while hanging from a twig. This behaviour is often observed in the field
as well.
All the tubes were held in a large glass container, with a relative humidity of
about 75%, and about the same air temperature as the laboratory. The widely
varying surface temperatures measured in very thin vegetation do not usually occur
in laboratory rearing, and as a consequence the development of the nymphs is
retarded as compared to that in the field. Because these observations were required
before work in the field could begin, in 1953, field sampling could not be started
B. M. LENSINK : Distributional ecology of Acrididae 377
before the season was half over, causing us to miss the period from hatching to
the occurrence of the youngest nymphs.
Thorough observation of the various cultured instars yielded several characteris-
tics that were very useful for recognition of the insects in the field. The first and
second instars are easily distinguished from those of the third and fourth instars:
the latter instars show a distinct beginning of the development of the wings, which
during the fourth instar extend past the second tergite of the abdomen. The first
and second instars are more difficult to distinguish from each other. The first
instar of the three species is generally smaller than the second instar. In addition,
the pronotum is shorter than the head in the first instar and about as long as the
head in the second. The second instar shows a broadening of the posterior edges
of the mesonotum and metanotum, and this characteristic can be easily observed.
In distinguishing the corresponding instars of the three species, consistent use
was made of the general shape of the antennae, the darker lateral borders on the
back part of the abdomen (linea lateralis) and the little lines on the edges of the
dorsal side of the pronotum (carina lateralis) (CLARK, 1943).
M. maculatus is the easiest to distinguish from the two Chorthippus species
because from the first instar on, this species has easily recognizable antennal clubs.
In addition, the linea lateralis does not form an unbroken line but per segment
forms an angle with respect to the dorsal abdominal line.
Chorthippus species do not have clubbed antennae. In the first instar of C. br-
guttulus, the linea lateralis is broken by a per segment shift. This also appears
in the first instar of C. brunneus, but in the same nymphs of C. albomarginatus
these stripes run almost continuously from front to back. The second, third, and
fourth instars of C. biguttulus are distinguished from the same instars of C. albo-
marginatus by the fact that the carina lateralis of the pronotum of the former is
not straight, but bent inward.
THE NUMBER OF INSTARS
For M. maculatus, C. biguttulus, and C. albomarginatus four instars and the
adult are distinguished in the field. UVAROV (1928) reports that the most of the
species of Acrididae have five instars, while BEI-BIENKO (quoted by Uvarov,
1928) has observed four instars in various grasshoppers occurring in Siberia.
RICHARDS & WALOFF (1954) distinguish four instars in the field. BEIER (1956)
states that the nymphs go through five or six moults; here the moulting of the
‘’vermiform’’ larvae is probably included. When higher mean temperatures occur
during development, one moulting can be omitted, and female nymphs often have
one more moult than the males (BEIER, 1956).
During rearing of the nymphs from the egg to the adult stage in the laboratory,
more than four instars were found in a few C. biguttulus. Two of these insects
moulted, after passing through the first instar, to small grasshoppers whose
characteristics, as used in field-identification, were not distinguishable from the
first stage. Another nymph passed through the first instar twice and through the
second instar twice during its development. The subsequent development of these
insects agreed with that of the other nymphs of the same species. In establishing
378 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
the number of instars, the vermiform larval stage preceding the first instar was
not included.
THE SAMPLING TECHNIQUE
In preparing the distribution study of the three species of grasshoppers, it was
my intention to become familiar with the distribution of the individual immature
stages and the adults in the various types of vegetation. The technique had to be
adapted to the collection of the required insect material and be related to the
behaviour of the insects. The older nymph instars and the adults move with great
rapidity, particularly in sunny, dry weather, while the younger nymph instars show
a tendency to remain concealed in the vegetation.
Other attempts have been made, in various ways, to determine the size of a
grasshopper population as representatively as possible, e.g., by the male song; by
catching, marking and releasing the insects (LINCOLN Index); by beating strips of
vegetation and attempting to count and identify the escaping insects; and by repeat-
ing a fixed number of passages with a net through the vegetation. All these
techniques, however, provide a more or less inadequate picture. Often the nymphs
are excluded from consideration. The technique of catching, marking and releasing
is only suitable for adults because during moulting the nymphs lose their external
markings. The beating of strips of vegetation is only applicable in identical types
of vegetation and strips of appreciable size (BALOGH, 1958). The yield is also
highly dependent on weather conditions and the moistness of the vegetation.
Taking into account the nature of the insects and the vulnerability of the
nymphs, a technique was developed for this investigation that served quite satis-
factorily under the given conditions, although it was somewhat laborious. The
requirements set for this sampling technique are summarized in the following
six points:
(1) All adult insects and nymphs must be caught without injury and later
released;
(2) During sampling no immigration or emigration may take place in the
quadrat;
(3) The vegetation must be disturbed as little as possible during the sampling;
(4) The apparatus used for sampling must be easy to manipulate;
(5) The sampling must not require too much time;
(6) Sampling must be feasible over long periods without loss of accuracy.
These considerations led to the construction of the “box, an apparatus con-
sisting of four wooden frames, each 75 cm high and 175 cm long, on which gauze
is stretched. The surface to be covered in this way is about three sq.m.
The grasshoppers were caught in a glass tube, identified, and held away from
sunlight. After all animals were caught, the insects were released within the
sampled area. This technique consumed the most time during the spring, when
the vegetation is populated with large numbers of nymphs. Under such conditions
the sampling of the 50 quadrats sometimes took as much as three days. Later in
the season they were worked in a day and a half.
B. M. LENSINK : Distributional ecology of Acrididae 379
The box is difficult for one person to operate alone. Collection of data was
consistently done by two persons, one of whom found the insects while the other
checked the sides of the box and recorded the data. Working together, the two
could move the box to a new site quickly and easily. Since the terrain chosen for
the investigation lies around the Biological Station, no difficulties were encountered
with the transportation of the box.
The box was placed over permanent quadrats. Inside it a small crate from which
the grasshoppers could be hunted, was placed over the vegetation. By moving the
crate the vegetation could be searched from various spots. Only the edges of the
crate pressed against the vegetation, keeping damage at a minimum. The vegetation
was searched thoroughly, which, especially in the thick grass cover, took a great
deal of time.
During the early summer when the field population was mainly nymphs,
sampling was done every week when possible, the period later being lengthened
to 14 days. The times of day and weather conditions suitable for sampling are
important. This work cannot be done at all times and under all conditions. When
the vegetation is wet (e.g., in the early hours of the morning in the dense grass
vegetations and after rain) and the small nymphs are disturbed, they come into
contact with the droplets of water; observations have shown that many of them
do not survive this contact. All sampling was therefore done in quiet, dry weather
in dry vegetation, usually between 11:00 p.m. and 5:00 a.m. When the wind
increased in velocity during sampling, the box was anchored with guy-ropes.
During the three years in which the study was carried out, 50 permanent quad-
rats each measuring three square metres were sampled with the box, 17 times
in 1953 and 1954, and four times in 1955. In addition, in 1955 another 23
quadrats were chosen and sampled 17 times in order to increase the number of
observations at specific places.
In 1954, a small box measuring one sq.m and 20 cm in height was used to
sample 55 quadrats in the period during which the first instar occurred in the
field, in order to provide more information concerning their hatching sites.
The systematic sampling of the quadrats was accompanied by a daily visit to the
field for observations concerning the occurrence of the three species outside the
quadrats. The collection of grasshoppers for laboratory examination was not done
on the terrain of investigation, but elsewhere in the dry grasslands of the landward
dunes.
PERIODICITY AND PHENOLOGICAL DATA
This chapter treats the periodicity associated with the grasshopper and, where
possible, its relationship with a number of climatic factors.
In winter, no active stages of the three species are seen in the field. Only the
eggs survive the winter. During April and May, large numbers of the first instar
appear. Within a month and a half the nymphs develop into adults, and the latter
populate the field during the summer and part of the autumn. In this period, the
eggs are laid.
Although it was not always possible to determine the time of the appearance
380 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
and disappearance of each instar with equal exactness, the field observations give
an adequate picture of the occurrence of these stages in the course of the year.
The adults were recorded according to sex.
A daily search for the first nymphs was made from 1 April on, in order to
determine the commencement of the activities in the field. The date on which the
first instar of M. maculatus appeared could be determined exactly because this
was the first species to be observed in the field. The first nymphs of the three
species appeared in 1953—1955 in the last ten days of April and the first ten days
of May (Table 3).
Table 3. Occurrence of the first nymphs of M. maculatus, C. biguttulus, and C. albomargi-
natus in the field in 1953—1955.
1953 1954 1955
M. maculatus 23 April 21 April 23 April
C. biguttulus 5 May 6 May 29 April
C. albomarginatus 8 May 6 May 8 May
Wide variations in the appearance of the nymphs do not occur in the successive
years in spite of the differences in weather conditions in the periods preceding
hatching. The monthly reports on general weather conditions issued by the
K.N.M.I. (Royal Netherlands Meteorological Institute) at De Bilt give the fol-
lowing data: in 1953, January and February were rather cold and clouded winter
months; March and April followed with dry weather and a more or less normal
temperature curve. In 1954, the first two months of the year were again cold, but
March was warmer than normal and April sunny, dry but rather cold. In 1955,
January, February, and March were cold, April was dry with normal temperatures;
March and April were rather sunny. Two factors predominate in these reports:
temperature and sunshine.
According to RICHARDS & WALOFF (1954), the hatching of the eggs in the
field depends upon the air temperature in the spring months. The average tem-
perature of the month in which the first eggs hatch is ordinarily 10° C or higher
(8.3° C in 1950). In the cold spring of 1951 the average temperature in April
and May was 6.7° C and 9.4° C respectively. In that year in Silwood Park, Berk-
shire (England) the first nymphs of five species (including M. maculatus) under
study did not appear until the first weeks of June, six weeks later than in 1948
and 1949.
No air temperatures are known for Voorne in 1953—1955. The nearest mete-
orological station is in Hoek van Holland, which did not publish continuous ob-
servations. Although the station in Vlissingen is far away from Voorne, the
average temperatures recorded there do not vary greatly from those on Voorne
(STOUTJESDIJK, oral communication). In addition, the island lies in a climatologic-
al area with mild summers and very mild winters, to which Vlissingen also
belongs (MÖRZER BRUIJNS & WESTHOFF, 1951).
If a comparison is made with the values measured in 1954 at the Vlissingen
station, then the eggs of M. maculatus and C. biguttulus hatch in the field at an
B. M. LENSINK : Distributional ecology of Acrididae 381
Table á. The average air temperature in March and April, and the first ten days of May
and the time at which the nymphs appeared.
March April | 1—10 May
1953 4,8° C STORIE STAGE
M. maculatus C. biguttulus
C. albomarginatus
M. maculatus C. albomarginatus
C. biguttulus
|
1954 | GMC OTA IAC
M. maculatus C. biguttulus
C. albomarginatus
1955 254€ TE SIE
average air temperature of 6.5° C and those of C. albomarginatus at + 10° C or
higher (Table 4).
After a very cold March in 1955, the average month’s temperature of April
reached a normal value and the grasshoppers appeared on dates which do not
differ greatly from those of the preceding years. Low air temperature apparently
has no influence, in the direct sense, on the rate of the eggs’ development. The
amount of sunshine and the amount of radiation heat dependent on it are more
important because they determine the course of the temperatures in the surface
layer of the soil. The radiation heat was not locally measured, but it may be as-
sumed from the above-normal amount of sunshine ın March, 1955 (46%, as
against 33% and 24% in 1953 and 1954, respectively) that the eggs received so
much heat that their development was not retarded in this cold month.
The first instar nymphs are seen in the field during a period which is longer
than the time required by the instar for its development. In 1955 this period was
79 days for M. maculatus, 73 days for C. biguttulus, and 51 days for C. albo-
marginatus. If the minimum developmental time of the first instar is considered
to be ten days, then nymphs must have hatched from the eggs during periods of
69, 63, and 41 days for the respective species.
Table 5. The numbers of the first instar of M. maculatus observed in various types of
vegetation on 8, 8, and 4 sq.m in 1954 (see also Table 11).
Type I | II and II IV
21.IV 2 0 0
23.1V 10 1 0
25.1V 46 0 0
27.1V 109 20 0
29.1V 148 32 0
1.V 326 83 6
3.V 102 108 115)
5.V 150 85 9
382 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
In this connection the place where the egg is laid is unquestionably of im-
portance. For its development, the egg requires a given amount of heat which can
be expressed in "day degrees” (e.g., ALLEE et al., 1950). One “day degree” is
realized if the average temperature during 1 day rises 1 degree above the develop-
mental zero point. This quantity will be realized more slowly in places where the
soil is covered than where the soil can profit maximally from solar radiation. For
the investigation into the hatching sites (1954), the hatching of larvae of M.
maculatus was observed from 21 April on (Table 5) in the very thin vegetations
in which part of the soil was not covered with moss or herbs. In vegetation of
types II and III, in which the moss cover extended over the entire surface, the
hatching began between 25 and 27 April, while in type IV, a vegetation with a
carpetting layer above the moss cover, the first newly-hatched larvae were seen
about 1 May.
RICHARDS & WALOFF (1954) report similar findings. They indicate still another
factor influencing hatching time, namely the date on which the egg was laid.
Observations in C. parallelus and C. brunneus point in this direction.
Unfortunately, the results of the winter survival of egg pods collected by me on
various dates are so unfavourable that no conclusions can be drawn from them.
When large numbers of nymphs of all instars populate the field later in the
early summer, a newly-hatched larva is rarely found. On 4 June, 1954, for instance,
on 150 sq.m 27 first instar nymphs were found, and on 23 June only one; on
22 June, 1955, on 69 sq.m 25 nymphs and on 11 July only three. This indicates
that during the last twenty days on which first instar nymphs occur in the field,
the numbers are very small, and thus that most of the nymphs have hatched in a
period of a month after the appearance of the first.
In 1953 and 1954, first instar nymphs of M. maculatus and C. biguttulus were
again seen in the field later in the season, appearing about a month after the first
group had disappeared. In 1955 a third instar nymph of M. maculatus was found
on 6 September after the first group had disappeared by the end of July. For
C. albomarginatus a newly-hatched larva was seen only once, on 4 August, 1953,
half a month after the disappearance of this stage. Certainly these animals are of
a second generation of that year.
We were not succesful in raising a second generation within one season in the
laboratory. Egg pods collected in the summer months and held at room temperature
in damp sand did not hatch in the autumn or later. If they were collected from the
field after the winter, for instance in February, they hatched in large numbers.
RICHARDS & WALOFF (1954) report the same experience. Eggs of three species
of grasshoppers studied by them continued to develop for a while after laying and
then entered the diapause. Towards the end of November, if the eggs were brought
into damp filter paper at a temperature of 25° C larvae hatched after some time.
Eggs of C. brunneus laid at the end of August and transferred immediately to a
temperature of 25° C failed to hatch. These authors were able, however, to raise
generations of grasshoppers throughout the year by giving eggs a cold treatment
in which temperature and duration of treatment were important.
Many insects in diapause die without development or grow in an irregular man-
ner when they are brought into temperatures which might be expected to promote
B. M. LENSINK : Distributional ecology of Acrididae 383
Table 6. The duration of development of the nymphal stages and the date on which the
adults disappeared from the field, determined for M. maculatus, C. biguttulus, and
C. albomarginatus.
| Year Period Duration Last observed
1953 23.1V—30.V 27.X
M. maculatus 1954 21.IV— 3.VI 44 ,, | 27.X
1955 23.1V—20.VI Sne | 1.XI
1953 5.V — 1.VII SSL 1.XI
C. biguttulus 1954 6.V —29.VI SOS 1.XI
1955 29.1V—11.VII JEU 1.XI
1953 8.V —20.VI AD 16.X
C. albomarginatus 1954 6.V —14.VI 40 ,, 15.X
1955 8.V — 5.VII SON. 20.X
development (LEES, 1955). The diapause is apparently interrupted when the grass-
hoppers are exposed to a low temperature for some time. In this connection it is
not clear, in the absence of further investigation, how the occurrence of a second
generation in the field in the summer is to be explained.
The succession of the nymph stages occurs rapidly. The dates on which the first
and the last nymphs of each stage are found in the field were not definitely
determined, but since the date on which the first adults appear could be determined
exactly, the total duration of the development of the nymphs can be calculated
(Table 6).
Table 7. The numbers of males and females found on various dates in 1954 on 150 sq.m.
M. maculatus C. biguttulus C. albomarginatus
4 2 ö 2 3 ©
cre, li di nl RES
25.V 0 0 0 0 0 0
4.VI 28 5 0 0 0 0
14.VI 66 25 0 0 4 0
23.VI 98 47 0 0 7 0
29.VI 107 49 1 0 18 2
9.VII 111 61 0 0 54 11
20.VII 116 107 1 0 97 38
29.VII 93 89 3 3 48 39
6.VIII Sil 75 | 11 4 35 31
14.VIII 30 53 10 7 26 30
28.VIII 23 41 | 16 16 18 24
7.IX 21 34 11 23 7 15
28.IX 4 5 9 9 1 3
DK 3 4 2 6 1 2
The duration of development of the nymph stages of these three species was
appreciably longer in 1955 than in the two preceding years. The month of May,
1955, was cold and wet, June was on the cold side with a normal amount of sun-
384 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
shine. Apparently in May, 1955, the insects had too little opportuntiy to sun them-
selves.
As soon as the adults appeared in the field a distinction was made between the
sexes. Table 7 gives the numbers of males and females found in the series
sampling on 150 sq.m.
The males of all three species were found in the field earlier than the females.
Initially, the numbers of males were higher, but in the month of July for M.
maculatus, and in August for the other two species, they were overtaken by the
females.
Thereafter, towards the end of the season, more females than males were found.
The numerical variations and the moment at which the largest numbers were found
show distinct differences. The males and the females appear in the field at some-
what different periods, the nymph stage of the females apparently having a longer
period of development than the males. It is unlikely that the eggs with male insects
hatch earlier than the eggs with females.
RICHARDS & WALOFF (1954) found for C. brunneus nymphs a sex ratio of
1:1, and a surplus of males among the adults. For C. parallelus and Stenobothrus
lineatus, however, they found more females among the nymphs and more males
among the adults. Although their figures for M. maculatus are rather small
compared with those for the other species, there appear to be more males in the
first stage, equal numbers in the second, and a surplus of females in the other
stages.
Although I collected no data on the sex-ratio of the nymphs, it appears that
if the sex is determined per sample for the adults, the ratio during the season is
first in favour of the males, and later in favour of the females. C. biguttulus
shows the most irregular fluctuations, probably as a result of the high activity of
this species in the terrain under study and its preference for the borders and the
low shrub which could not be included in the sampling. It is for this reason that
the numbers on the sampled surface are consistently small.
The adults remain in the field until late in the autumn. After September the
numbers decrease rapidly and the insects disappear entirely towards the end of
October and beginning of November when the effect of the night frost becomes
noticeable throughout the field (Table 6). In 1953 and 1954, C. albomarginatus
disappeared half a month earlier than the other two species. In 1955, this species
was not observed in the dense vegetation after 10 October and in the other
vegetations after 20 October. Especially in vegetation of type IV, the microclimate
in the autumn is appreciably less favourable for the grasshopper, which requires
warmth, than in the less dense growth because in the former on clear nights the
night temperatures fall below zero very early in the season.
THE NUMBERS OF GRASSHOPPERS
The sampling of a large number of quadrats at various times during the season
provided information concerning the numbers of insects present in the quadrats.
It was not intended to use these figures for population studies and for drawing
conclusions about such factors as the birth-rate and mortality of the total population
B. M. LENSINK : Distributional ecology of Acrididae 385
in the area under study. Inventories made at regular intervals were intended to
provide information about the composition and the numbers of the grasshopper
population in a given type of vegetation.
The collected data formed a picture of the following points:
(a) the numbers of insects observed in this terrain in the three years of the
investigation;
(b) the way in which the numbers of the nymphs and adults vary in the course
of the season; this point is of particular importance for evaluating whether or not
the field sampling was adequate;
(c) the composition of the population at various times in the season.
In 1953, 1954, and 1955, from the beginning of the hatching of the nymphs
in the field until the disappearance of the insects at the end of October, samples
were taken 17 times on suitable days. In the respective years, 150, 150, and 69
sq.m of the field were inventoried. Tables 8, 9, and 10 give, for each date of
sampling, the numbers of nymphs of the various stages and of the adults for these
three years.
The highest density of the three species was observed early in the season, namely
at the end of May or beginning of June, the period in which nymphs are mainly
seen. M. maculatus was the most numerous species. At the time of peak density,
on quadrats of types I—V in which this species was found predominantly, we
found 12.3 insects per sq.m sampled in 1954 and 13.8 insects in 1955. The next
most frequent species was C. albomarginatus, on type HI—VI quadrats giving 5.0
insects per sq.m in 1954 and 3.8 in 1955. The least frequent species was C. bigut-
tulus in types I—VI, with 1.4 in 1954 and 0.8 in 1955.
Table 8. Numbers of nymphs and adults (a) on the 50 quadrats sampled in 1953.*
M. maculatus C. biguttulus C. albomarginatus
Instar 1 2 3 4 a 1 2 3 4 a 1 2 3 4 a
22.VI 12.857 10971237 742 SR 3820172 6 — 48 62 102 30 3
4.VII ORO SE OI 23 NAS I 5 1 Mh je Vr EE)
11.VII — 2,1 SO VI We 7023026512. 2 eI Osh ES)
16.VII MEZ COS Tio De BD 10050160400 75
28.VII aa MOST A Ton ZON Tl Mery SO IG MTBE
4.VHI DA a e e aN En RL MENT ENGE
11.VIII ZA II O EG eo aye EG
18. VIII SEE Gal TANT A ES AE NE 3
26.VIII ee OR AG N SO)
LIX add fB erp
8.IX a Sa ee a 32
15.IX as ee en en 22
24.IX I E O E e PAM CR IB
DINE ME E ZON OI I. 6
TX E N ET pi TE ANTE O EN eee
16.X A I | en ee PL ARE
27.X nn rs LL or Dr ER A no
* In this and the following tables containing data pertaining to the sampling of the quadrats,
the date indicates the time at which sampling was begun (see p. 378).
386 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Table 9. Numbers of nymphs and adults (a) on the 50 quadrats sampled in 1954.
M. maculatus | C. biguttulus | C. albomarginatus
Instar 1 2 3 4 a | 1 2 3 4 a | 1 2 3 4 a
AIV | 13 — — el ~ ~ —
6.V 501 16 — — — 1 — — — — 4 — — — —
25.V Sk G45 ZL 37 = 88 11 — — — | 360 183 © = =
4.VI DITA SAT LOENEN SSA 4 — — | 121 382 41 i =
14.VI 2 162 240 305 91 |107 87 11 D == 23 153 150 57 4
23.VI 1 53 166 229 145 36 120 25 4 — D 72 iS) OO 7
29.VI — 14 111 138 156 14 60 32 1 1 — 37 109 119 20
9.VII = Dr Di CE ye NP GO) Sythe} — 14 58 112 65
20.VII = = 70226675223 TWAS 33/0035 1 — — 11 95 135
29.VII 5 2 5) 2 U 1 O 19. AS 6 “lla 2237)
6.VIII 8 3 1 3 126 | — 1 020,15 =. = 2 5 66
14.VIII 9 SIM EN — 5 Di Ly = = = 3 X6
28.VIII 3 11 — — 64 — 2 — 522 1 42
7.1X = 4 9 20255 — 2 — — 34 — — — — 22
28.IX = 1 9 == = (IS, dn 4
12.X = = — 1 7 — = U == 8 oe 3
In the period in which the adults populate the field, numbers have become small
and density shows strong local variation. The largest number of adults of M.
maculatus found on one sq.m is 8.0, of C. albomarginatus 28.6, both in 1954.
The latter number was found on 20 July, 1954 in Quadrat 6 in a high, dense
vegetation of Calamagrostis epigejos.
RICHARDS & WALOFF (1954) noted, concerning C. albomarginatus, that the
largest number of adults found by them in England was only 2.0 to 2.2 insects
Table 10. Numbers of nymphs and adults (a) on the 23 quadrats sampled in 1955.
M. maculatus
Instar | 1 2 3 4
25.IV — — — —
29.1V 69 = =
8.V 402 26 1 —
20.V 566 122 10 —
30.V 5321870037 3
6.VI 15209232510 60017
16.VI 86 187 129 58
23.VI 26 169 143 101
27.VI 3 75 110 108
5.VII 2190165 0 41
11.VII SNS CIN CO
25.VII — — SM?
11.VIII — — — 1
23.VIII — — — 2
6.IX — — 1 —
3.X — — — —
C. biguttulus
C. albomarginatus
lcabou |
| 5 |
WO OW we Ke
Poi EN
As PE ENTER
US 1 SoS =
554 110005) MES
DP UE 1 — —
4 77 30 — —
B. M. LENSINK : Distributional ecology of Acrididae 387
per sq.m in the favourable year of 1949. In West Siberia, where this species some-
times occurs in such large numbers that the term plague is used, RUBTZOV (as
cited by RICHARDS & WALOFF, 1954) reports numbers in the range of 100 to 150
per sq.m.
In the course of the season the numbers of insects gradually decrease. Charac-
teristically, the curves representing the increase and decrease in the numbers of the
various stages of the insects are smooth. There are thus no deviating observations
suggesting great unreliability in the counts. As the season progresses we also see
a series of gradually-occurring peaks, each time lying lower, reflecting the numbers
of insects of the successive instars. This is in agreement with expectation, since
as a result of mortality each stage would be present in lesser numbers than the
preceding one.
The composition of the population on the total surface sampled can be followed
easily and offers good comparative material for the study of the data derived
from the separate types of vegetation.
THE HATCHING SITES
Introduction
The investigation of the places at which the larvae emerge from the eggs is
important because it is from these places that the active stages of the grasshoppers
make a choice among the possibilities offered by the vegetation. Information was
already available concerning the requirements governing the choice of oviposition
substrates by the females of the three species (LENSINK, 1962). This information
provides a useful indication of the places in which the initial appearance of the
insects can be expected.
The hatching sites were studied in 1954 by means of the quantitative sampling
of a large number of quadrats (55 in all) in various types of vegetation which
were visited every two or three days, including quadrats on horizontal ground as
well at the north and south sides of slopes.
The investigation had to be suspended a month after the appearance of the
first instar: by that time there were so many nymphs in the field that sampling
became unreliable. As a result of their increased activity, many of the larger
nymphs jumped over the edge of the box. The greatest quantity of data was
obtained for M. maculatus, because C. biguttulus and C. albomarginatus hatched
later. However, numerous observations done apart from the sampling, as well as
the data on the distribution of the first instar, were sufficient to provide a good
idea of the hatching sites of C. biguttulus and C. albomarginatus.
The hatching sites of M. maculatus
Table 11 gives the numbers of first instar of M. maculatus found on various
dates on the quadrats sampled, as well as the type to which the vegetation belonged
and also exposure, total vegetation coverage in per cents, the proportion of mosses
and herbs in these percentages and the height of the vegetation in centimetres.
388 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Table 11. The numbers of the first instar of M. maculatus on 55 quadrats measuring
mn
EEE
Total Herb Moss Bare |Height of
Quadrat | coverage layer layer surface |vegetation | 21.1V 23.1V SN:
in % in % in % in % in cm
"NW-NE-slopes” types I, II, and IV
1 100 80 30 0 10 = a È
F 100 60 50 0 5 “a oh 4
18 95 40 95 5 5 — _ A
ap 100 25 95 0 10 — = A
U 60 40 30 40 5 = ee a
Y 100 70 50 0 10 = Pie: a
BB 100 90 30 0 15 == 28 MI
NN 85 40 70 20 5 = me
”SE-SW-slopes” a) type I |
B 50 40 50 40 10 = 1 3
C 60 55 10 40 10 ae D 6
E 90 20 90 10 5 = = =
H 40 40 50 40 10 1 E =
9 50 40 40 60 10 1 2 7
J 70 25 70 30 10 = 1 =
K 60 40 50 40 10 6 16 29
HH 75 30 70 30 10 — = 4
DE 70 30 50 35 10 1 2 2
b) types II, III, and IV
A 100 80 30 0 15 = == 1
G 95 55 60 5 10 — — —
L 100 60 50 0 10 = 2 1
P 95 25 95 5 15 1 = =
V 90 70 30 10 10 = DI 1
GG 100 50 95 0 15 — == ="?
”Flat field” a) type I
M 95 60 60 10 15 — en 1
N 25 15 25 75 10 = — 3
O 30 25 10 15 10 — — 6
Q 30 25 30 70 5 2 10 32
W 80 60 40 20 20 a ze 2
KK 60 40 20 40 5 = — —
JJ 80 70 30 30 5 = — 1
S 95 50 70 5 5 — = 1
b) types II, III, and IV
2 90 75 30 15 20 = — =
R 100 85 50 0 10 — 1 =
31 90 80 70 10 10 = — =
Zi 95 75 60 10 10 = — à
CC 90 75 50 10 5 = = “à
MM 95 70 70 5 5 — a a
7 95 50 90 5 10 = — a
8 95 50 90 5 10 = = a
11 100 50 80 0 10 en — 1
EE 95 50 50 10 5 — =
FF 100 90 20 10 5 = vari
B. M. LENSINK : Distributional ecology of Acrididae 389
one sq.m, in various types of vegetation, between 21 April and 18 May, 1954.
mmm
|
27IV | 29.1V 1.V 3/4.V 5.V 7.V 10.V 13.V | 17/18.V
|
= = == 1 3 1 4 11 1
= == — — — — 4 11 5
= E 2 2 = 5 16 28 5
= 1 = — = 2 4 2 2
LA = 1 3 — 1 9 11 1
— = = = = 2 1 3 1
= — — — — — 1 14 5
= 14 13 3 8 9 26 8 19
5 6 14 12 11 18 16 14 8
— = — 2 4 3 5 25 4
1 4 0 3 6 13 36 33 8
7 11 12 8 10 35 15 19 3
6 4 9 4 9 17 9 10 15
49 49 22 3 7 DIL 16 8
-- 3 7 3 1 18 4 177;
16 6 15 1 2 20 6 11 5
— = 2 3 3 1 7 9 8
— = — D 12 18 16 6
6 4 1 8 8 14 13 8
16 20 7 12 22 18 12 15
1 3 1 2 5 10 7 11 1
— = — 5 1 3 12 7
17 21 18 24 24 36 44 32 21
3 32 5 28 28 23 38 17 14
24 34 13 19 19 46 60 59 11
66 80 17 22 22 55 52 37 6
27 197 22 27, My 72 72 65 20
2 24 7 7 7 23 23 12 9
7 13 10 8 8 37 23 15 15
5 10 15 15 38 49 28 6
= 1 1 1 3 8 21 9 8
9 13 23 29 26 35 23 5 5
7 9 30 25 19 13 57 38 13
2 3 7 25 11 49 79 129 26
= 1 3 13 8 14 37 49 9
2 3 15 11 16 42 12 22 10
= 2 2 3 — — 7 12 10
= — 2 1 2 1 2 31 18
n 3 = 7 19 20 5
4 5 11 sun 2 5 18 13 5
3 1 2 7 25 65 11
390 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Table 11, continued. The numbers of the first instar of M. maculatus on 55 quadrats measuring
Total | Herb Moss Bare | Height of
Quadrat | coverage | layer layer surface |vegetation | 21.IV 23.IV 25.1V
in % in % in % in % in cm
c) type V
D 90 19 25 10 15 = = =
5 100 100 10 0 15 = = =
10 100 85 60 0 15 — = —
DD 95 90 20 5 5 — = —
38 100 95 40 0 10 = = —
d) type VI
3 100 100 >) 0 15 == = =
4 80 80 10 10 40 => = —
6 100 100 0 0 30 = = =
30 100 100 0 0 15 = = =
16 100 100 15 115) 10 — — —
xX 100 100 0 0 15 Ta = —
14 100 100 0 0 15 = TT i
AA 100 100 5 5 10 — = =
The following may be derived from the data in this table.
The first nymphs of M. maculatus are found most densely in the areas indicated
by ”SE-SW-slopes” and "Flat field”. In these areas, the quadrats with type I
vegetation yielded large numbers. Those with vegetations of types IITIV gave
lower hatching figures. In the "Flat field” types I—VII are also found. The num-
bers of nymphs in type V are small. In type VI the figure was zero or very nearly
so. Type VII could not be systematically sampled with the box.
The first nymphs appear much later on the quadrats on the "NW-NE-slopes”,
in 1954 as much as ten days after the first nymphs had hatched.
In terms of the rate at which the number increased, it can be seen that for the
twelve samplings made between 21 April and 17—18 May in 1954 there was a
rapid increase in the population of the ”SE-SW-slopes” type I quadrats. Except
for lower initial rates, the ”Flat field” type I quadrats show almost the same
picture. For types II—IV, the rate of increase is slower. The slowest rate is found
in the "Flat field” types II—IV.
The amount of heat originating from sunlight is of course an obvious factor to
consider in relation to these phenomena. In this connection the figures for the
’NW-NE-slopes” become meaningful: the first nymphs appear on such slopes
after all the other quadrats are populated. The importance of solar radiation for
the rate of development of the eggs has already been discussed. In places with a
well-developed surface cover, the eggs receive less heat than in uncovered sandy
spots. In addition, in the spring appreciably less solar radiation reaches the north
slopes than the south slopes (STOUTJESDIJK, 1959), which explains the retardation
in the development of the eggs laid on the former. Although the field on the
north slopes lends itself well for oviposition, I have never been able to find newly-
hatched larvae on them, and this also holds for the skins of the moulting vermi-
form larvae which precede the first instar. This raises the question of whether we
B. M. LENSINK : Distributional ecology of Acrididae 391
one sq.m, in various types of vegetation, between 21 April and 18 May, 1954.
17/18.V
11 20 1 4
1 15 1 4
any 3
= — — — 2 1 1 1
— -- — 1 — 1 1 4
have to do here with migration from other localities. Several factors may be
involved. The insects may have left their hatching sites of their own volition,
some of them settling on the north slopes. Such migration would initially be small,
because the young are small and not very active. Migration could be promoted by
the passage of large animals and of people through the hatching areas; this can
be seen to cause the insects to jump away in all directions. The wind may give
direction to these movements, as can be observed when walking through vegetation
when the wind is blowing hard: the jumping grasshoppers are swept in one
direction by the wind. Before the first instar was found on the north slopes, a
moderate to strong west to south wind blew from 1 May, 1954 on, so that the
nymphs which hatched on the south slopes could have been blown into the north
slopes.
If we now return to the absolute figures given in the table for the various
sectors, we note large differences. In some places large numbers of nymphs are
found, in others very few. In a part of the field with sites suitable for oviposition,
few or no young nymphs were found, and in the summer months few adults were
seen in the same places. The oviposition substrate is evidently only one of the
factors to play a role in distribution.
In most places, the observations made in 1953 and 1955 agreed with those
made in 1954. In 1955, however, there were local modifications in the vegetation.
In the cold, damp spring of 1955, such grasses as Helictotrichon pubescens
reached their full growth very early and the structure of the vegetation in which
they occurred, showed marked changes earlier in the season than would normally
be expected. In the autumn of 1954, many eggs were found in the soil of a part
of the terrain with a thin vegetation (type IV), but in 1955 the same locality
showed a much denser vegetation at hatching time because of the rapid growth
of these grasses. Nevertheless, large numbers of nymphs hatched in this moderately
392 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
dense growth (type V). In agreement with expectation, almost no eggs were laid
in this part of the vegetation in the summer of 1955. It follows from this
observation that such deviations from the normal can only be explained when the
situation in the vegetation during the period of oviposition is known.
Lastly, it can be seen from the table that on 3—4 May the numbers of nymphs,
particularly on the very thinly covered plots, were markedly reduced. On 3 May
there was a severe hail storm, just after sampling was begun, the entire field being
strewn with hailstones. Sampling was resumed after the vegetation had dried, and
was continued on 4 May. The numbers of insects found on the very thinly-covered
quadrats were appreciably smaller than those on 1 May (Table 5). The hail took
a great toll of the first instar nymphs. In the denser vegetation little or no effect
was seen, the nymphs having been much better protected. On 5 May, many newly-
hatched larvae were again found on the very thinly-covered spots. Other in-
vestigators, including UVAROV (1928), CHOPARD (1938) and RICHARDS & Wa-
LOFF (1957) have observed the phenomenon of a sudden increase in hatching
after rain. I have observed this phenomenon, apart from this case of a hail storm,
once after rainfall.
The hatching sites of C. biguttulus
Because sampling of the hatching sites had to be limited, for practical reasons,
to a period of a month, few data could be collected for C. biguttulus, the species
which hatches the latest. The nymphs of this species did not appear until 6 May
in 1954, 15 days after the sampling had begun.
Simultaneously with the investigation of the hatching sites, the large-scale series
sampling of the three sq.m quadrats was begun. Although the series sampling was
done at long intervals in the hatching period, it nevertheless provided sufficient
data to create an impression of the places where the first instar appears in the field.
Table 12 gives the numbers of first instar nymphs of C. biguttulus found on 46
quadrats measuring three sq.m, classified according to the six types of vegetation,
from 24 April to 4 June, 1954. Table 13 gives the results of the 1955 sampling
of 23 quadrats measuring three sq.m, from 25 April to 16 June. Where possible,
the table for 1954 is supplemented with data from the sampling of the hatching
sites.
In no case do the numbers reach the high values found for M. maculatus. Just
as for M. maculatus, the date of first appearance in vegetations with incomplete
soil coverage (type I) falls earlier than in vegetation with a complete moss cover
and a well-developed herb layer (types II—V). This can be seen very clearly in
the table for 1955. In the absence of part of the sampling on 14 May, 1954,
Table 12 gives only a slight indication in this direction.
The places where the most first instar nymphs of C. biguttulus were found,
according to Tables 12 and 13, have vegetations belonging to types I—V, especially
in the very thin grass cover, but also in dense, herb-rich grass vegetation. The
numbers of nymphs found in the dense, high grass cover (type VI) are very small.
If we assume that the nymphs of this species could migrate only very short distan-
ces during the periods indicated in the tables, these samples of the first instar give
a good picture of the hatching sites. To make this picture as complete as possible,
B. M. LENSINK : Distributional ecology of Acrididae 393
Table 12. The numbers of the first instar of C. biguttulus on various quadrats measuring
three sq.m in six types of vegetation from 24 April to 4 June, 1954.
AMIN Tir
Quadrat | 24.IV 6.V | AE
Type I 2 — 1 11
11 = = 16
Type II 7 — = 10 2 1
Type III 34 — —
Type IV 5 — — —
Type V 3 = =
Type VI 6 — == se Le
45 = =
|
|
|
e not sampled
— no larvae
394 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Table 13. The numbers of the first instar of C. biguttulus on various quadrats measuring
three sq.m in five types of vegetation from 25 April to 16 June, 1955.
| Quadrat | 25.IV | 29.IV | 8.V | 20.V | 30.V | ui evi
Type I 2 — — — — 1 4
B — 1 4 7 1 = =
9 = 1 — 1 —
de — — — — 1 1 —
L — — — 7 6 2 8
31 — a —- —- 1 6 6
C = — = 1 1 4 6
Type II 7 — — — — — — 2
Type IV 5 — — — — — — —
H = == = = 5 1 =
J — — — — eo or
10 — = = — 2 5 1
8 —. = = 1 as = 1
A = = = = 2 1 1
Type V D CS = = = = 11 13
4 — — — = = 3 ==
N — — — — = 3 =
Type VI 3 — — — — — 2 —
6 EN zE = ae a Lis =
F au La pus sr Du, ie ce,
G A DEI Bie Dee En view
K — — — 2 — ts
M el enn en die cite Er
a study of the oviposition sites was made during the winter. The greatest numbers
of eggs were found in bare, sandy soil and in sandy soil with a moss cover. A few
egg pods were also found in soils with many roots in small, bare spots in the
dense, herb-rich vegetation.
Laboratory experiments concerning the choice of the oviposition substrate are in
good agreement with these findings (LENSINK, 1962). The insects were provided
with a number of substrates, collected from the field and kept as much intact as
possible, representing bare sandy soil, sandy soil with a moss cover, dead grass,
and a grass tussock. The eggs were laid in the bare sandy soil and in the sandy
soil covered with moss, using the places in the latter exposed by splitting of the
moss. Oviposition also appeared possible beside the grass tussock where the soil
was well-supplied with roots. No eggs were found in the grass tussock proper.
C. biguttulus therefore appears to have very ample opportunity for oviposition:
its substrate is present throughout the field, only the very dense grass cover being
unsuitable.
The absolute numbers of nymphs found on various plots of each type of vege-
tation show wide variation, however. The position of the quadrat in relation to its
surroundings appears to be important. In this sense we find in type I, which shows
strikingly few adults during the summer, the largest numbers on Quadrats 2, 11,
29, and 31. With the exception of Quadrat 29, these quadrats lie in the immediate
B. M. LENSINK : Distributional ecology of Acrididae 395
Table 14. The numbers of the first instar of C. albomarginatus on various quadrats
measuring three sq.m in six types of vegetation between 24 April and 4 June, 1954.
| Quadrat 24. IV | 6.V | 14.V | 25.V ANI
ho
Type I 2 = — —
Type II U = = + 3 3e
Type III 34 = sedi
Type IV 5 — — 15 82 33
Type V 3 = — 15 10 1
Type VI 6 - = 2
Lox
CERRI Rs
Ho
00 A 00 MO \O LA NS À
|
|
e not sampled
— no larvae
396 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
vicinity of type V (dense, herb-rich vegetation) and type VII (Salix repens shrub),
vegetations in which many adults were seen. The other quadrats of type I are
distributed in the field among type II vegetation. This indicates that the
combination of environments suitable to the adult insect and a suitable oviposition
substrate plays an important role in the distribution of these insects.
The hatching sites of C. albomarginatus
For C. albomarginatus, too, the hatching site samplings provided little data
concerning the places where the first nymphs appear in the field. The nymphs of
this species appeared in 1954 on 6 May in the field, i.e., halfway through the
period of sampling. As for C. biguttulus, the data from the sampling of the three
sq.m quadrats in 1954 and 1955 give a rather good impression of the hatching
sites. Supplementary data is provided by the otherwise scarce data obtained from
the study of the oviposition substrate.
Table 14 gives the numbers of the first instar of C. albomarginatus found on
46 quadrats measuring three sq.m, classified according to the six types of vege-
tation, between 24 April and 4 June, 1954. Table 15 shows the results of the
large-scale series sampling in 1955 on 23 quadrats measuring three sq.m, between
25 April and 16 June. The numbers of the first instar found in the field are,
with a few exceptions, not very large. With respect to population numbers, C.
albomarginatus falls between M. maculatus and C. biguttulus. On only one spot
in the field did this species take the leading position, especially in 1953 and 1954.
The relevant quadrats were, in 1954 Quadrats 5 and 6, and in 1955 Quadrats 5,
H, and J.
The first instar nymphs of C. albomarginatus were found, according to both
tables, in all the types of vegetation, but the highest numbers were found in the
more dense vegetation, in 1954 in types II, IV, V, and especially VI, and in
1955 only in type IV.
The absence of the hatching of nymphs in type VI in 1955 is even more remark-
able in relation to the data from the summer of 1954, when large numbers of
adults of this species were found in this type of vegetation. In Quadrat 6, for
instance, situated in a part of the terrain in which large numbers of C. albomar-
ginatus were consistently found, gave on 20 July, 1954 the maximum number of
86 adults insects on three sq.m.
We can be certain that eggs were laid in the summer of 1954 in the dense, high
grass vegetations because they were found in a number of places between dead
grass blades. It is probable that there was a high mortality among the egg pods in
this dense vegetation, possibly caused by a deviation in the normal spring tem-
peratures. In the spring of 1955, the minimum values for the air temperatures
were often very low. Especially in the high, dense grass vegetation, temperatures
in the latter part of the night may reach very low values, with local frost. Further
investigation will be required to determine whether eggs can be damaged by low
temperatures in the last stage of their development.
The search for egg pods in the winter months showed that they had been
deposited not only in type VI vegetation, but also in the moss layer of types III
B. M. LENSINK : Distributional ecology of Acrididae 397
Table 15. The numbers of the first instar of C. albomarginatus on various quadrats measuring
three sq.m in five types of vegetation between 25 April and 16 June, 1955.
Quadrat 25.IV | 29.IV | 8.V | Bat Nvt
Type I 2 2 i au 5 2 3 1
B ze en er 2 3 = se
9 = = ae = 5 de =
11 = = _ 3 5 2 =
L sc e PI en En SE 2
31 = = = 1 1 ac =
€ se = ek = 2 Be 2
Type II 7 — — = — 6 3 1
Type IV 5 — = — 13 13 7
H au — 3 37 10 8
ji pr 2 1 24 17 7
10 — — ei 2 2 =
8 en en Lie = 2 Be, 1
A ze on a _ 3 2 en
Type V D au an zij 1 2 le 1
4 a = en Pai Sen er LL
N Dil pa er om 4 es DA
Type VI 3 = = — — — 1 —
6 Es eal ze al Bei, a, kas
E es in ar Aa er a ai
G 2 = = = 1 2 a
K uh DA x 1 2 LES as
M Es en at = 1 En di
and IV in the immediate vicinity of high, dense grass growths in which the adults
occur during the summer months.
Experiments concerning the choice of oviposition substrate (LENSINK, 1962)
support these data. C. albomarginatus was provided with a number of substrates
collected as nearly intact as possible from the field. From the substrates provided
(bare sandy soil, moss-covered sandy soil, dead grass, and a grass tussock) the
females chose not only the tussock and the dead grass but also the moss-covered
sandy soil. The eggs were laid against the stalks of the grasses and dead grass
leaves and between the moss plants. No eggs were found in or on the bare sandy
soil.
C. albomarginatus thus has a great many possibilities for the deposition of eggs
in the terrain in which the study was made, since its preferred substrates occur
widely. The wide variations in the numbers of nymphs on the various quadrats
within the types of vegetation on which the largest numbers occurred indicate,
however, that for this species, too, the oviposition substrate is only one of the
factors which determine distribution in the field. In 1954 in type III, the largest
numbers were recorded for Quadrats 39 and 44, in type IV on 5 and 38, and in
1955 in type IV on 5, H, and J. The largest numbers of the first instar were found
on those plots lying closest to the type VI vegetations where the most adults of
this species were found during the summer months. These plots lie at a maximum
398 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
distance of five metres of the edge of the high, dense grass cover. The remaining
quadrats in types III and IV lie at a great distance, in various parts of the terrain.
Within type I, too, Quadrats 11 and 31 lie within five metres of the dense,
high grass vegetations. The small numbers of nymphs occurring on these very
thinly-covered spots indicate the unsuitability of these vegetations for oviposition.
THE DISTRIBUTION OF THE GRASSHOPPERS DURING THE SEASON
The distribution of M. maculatus
The investigation into the hatching sites of M. maculatus shows that most of
the nymphs hatch in particular types of vegetation. During their development the
insects spread out over a large part of the terrain. This dispersion was followed
by means of a series of 17 samplings taken during the season in 1953, 1954, and
1955 in six types of vegetation. In 1953, after a preparatory period, sampling was
not begun until 26 June, when adults were already present in the field. The data
from these samplings are therefore not suitable for systematic analysis, but have
been used, where relevant, in the discussion of the results of the 1954 and 1955
sampling.
Comparison of the total number of nymphs of each of the four stages with each
other and with the adult stage occurring on the quadrats of each type of vegetation,
is sufficient to give a general picture of what takes place in the field. Tables 16
and 17 give these figures for the nymphs and the adults in 1954 and 1955.
In Table 16 the columns for the first and second instars have been corrected
because a change in weather conditions interrupted the sampling on 14 May. Only
22 of the 46 quadrats could be sampled. The loss of part of the sample, just in a
period when the nymphs of the first instar are numerous in the field, meant the
loss of a number of data. In order to provide comparable values for the table, an
estimate was made of the numbers of nymphs of the first and second instar which
could be expected on 14 May on the 46 quadrats. For this use was made of the
data collected from the 22 quadrats over the entire season. We then compared the
numbers of nymphs of the first and second instar found on 14 May and the num-
bers of nymphs of the first and second instar found on those 22 quadrats during
all samplings. This gave the part taken by the 14 May sampling in the total series.
Table 18 gives these figures.
It appears that the numbers of nymphs of the first instar found on 14 May on
22 quadrats in vegetation types I—IV (there were still almost no nymphs present
in types V and VI) represent almost half in type I and appreciably more than half
in the other types of the numbers observed with the entire sampling. Table 16
gives the estimated data for 14 May, arrived at by taking the total numbers of
nymphs of the first and second instar found during the other samplings and
multiplying by 2 and 114 respectively.
Tables 16 and 17 give the data for a number of quadrats in type I vegetation
separately, since in connection with their special position these quadrats show
marked differences from the others. For the same reason the data from the three
quadrats in type VI are given separately in Table 16.
B. M. LENSINK : Distributional ecology of Acrididae 399
Table 16. The total numbers of nymphs and adults (a) of M. maculatus found on the
quadrats of each type of vegetation during the entire season in 1954.
No. of quadrats
Instar 1 sampled
a —
Type I 430 115 5
Type I 121 146 2
Type II 200 101 3
Type III 400 560 8
Type IV 300 387 330 238 9
Type V 21 159 281 332 460 8
Type VI 1 15 26 43 45 8
Type VI 0 14 33 90 97 3
No. of samplings 7 8 9 9 13
Table 17. Same observations as Table 16, but in 1955.
No. of quadrats
Instar sampled
Type I 4
Type I 5)
Type II 1
Type II er Pa as a ar dh
Type IV 332 359 266 158 92 6
Type V 96 80 95 104 91 3
Type VI 0 6 8 10 11 6
No. of samplings 11 10 10 10 10
The data in these tables, taken together, give an idea of events in the field, but
three factors must be kept in mind:
(a) the numbers of quadrats within each type of vegetation are not the same,
so that the horizontal columns may not be directly compared (for instance, type II
is poorly represented in this sense) ;
(b) during the presence of each stage in the field the quadrats were not
sampled the same number of times, so that direct comparison of the vertical
columns is not possible (Tables 16 and 17);
(c) the intervals between two successive samplings are not always the same
(Tables 19 and 20).
It is clear that the nymphs and adults populate almost all the various types of
vegetation in the terrain. Small numbers are found only for type VI, the high,
dense grass cover. Within types I and II, it is striking that the number of nymphs
decreases strongly from the first to the fourth instar. In 1954 in type I the figures
for the first to the fourth instar averaged 12.3, 2.9, 0.9, and 0.6 nymphs res-
pectively per quadrat per sampling. In 1955 the figures were 20.4, 8.2, 2.6,
and 1.4.
This sharp decrease is not found for types III and IV. Here the average num-
400 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Table 18. M. maculatus, sampling, 14 March, 1954. See text, p. 398.
Type : | II Ill | IV
Instar 1 2 | 1 2 1 2 | 1 2
22 quadrats, 14.V
22 quadrats
all samplings, 1954 282 209 230 91 Dy 41
bers of nymphs per quadrat and per sampling for 1954 is 7, 9, 5, and 3 in type III,
and 5.3, 6, 4.6, and 3.4 in type IV; for 1955: 5, 6, 4.4, and 2.6 in type IV.
For the quadrats within type V the average values for 1954 are 0.38, 2.5, 4,
and 4.5, and for 1955: 3, 2.7, 3.2, and 3.46 nymphs.
A decrease in numbers is to be expected in all cases as a result of mortality
among the insects. Unfavourable conditions in a given environment will increase
this mortality. But unfavourable conditions in a given environment may also lead
to migration. The character of the first two types of vegetation suggests that in
the course of the early summer, climatic conditions are unusually unfavourable
for the grasshoppers. The vegetation supplies a distinct indication for this. It is an
obvious assumption that migration will occur from places where conditions are
becoming less favourable toward more protected vegetation. This assumption is
supported by the variations in the values for types III through V. In types III and
IV the initial decrease is zero, but it is later smaller than that in the first two types.
Type V even shows a distinct initial increase in numbers.
The discussion which follows concerns the course of the development in space
and time, within each type on the individual quadrats.
Type I. Vegetation of type I is characterized by the very thin structure of the
plant growth. Tables 19 and 20 give the numbers of the various nymph stages
and adults (a) found on quadrats belonging to type I in the course of the season
in 1954 and 1955, respectively. Many nymphs of the first instar were found on
these quadrats and during the first month after hatching this quadrats are the most
densely populated. On one quadrat on a south slope with a very thin cover and
extreme exposure to wind and rain, however, very few grasshoppers were seen
during the entire season.
During the early summer, the numbers of nymphs of the older instars decreased
rapidly on the type I quadrats in 1954 (Table 19) and 1955 (Table 20). Starting
with the third and fourth instars, the numbers are small compared to those for
the first and second instars. Adults are scarce during the summer, but in late
summer and autumn the numbers show a relative increase in comparison to the
other types. In the latter period the density of the adults has already fallen far
below the highest values.
On Quadrats 2 and 11 in 1954 and 1955 the situation took a different course.
The numbers of first to fourth instar nymphs did decrease, but to a much smaller
extent than in the other quadrats of this type. Nymphs and especially adults were
found on Quadrats 2 and 11 throughout the season. If the vegetation of these
401
B. M. LENSINK : Distributional ecology of Acrididae
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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
402
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B. M. LENSINK: Distributional ecology of Acrididae 403
two quadrats is compared with that of the others, little difference is found, but
there is a difference in their location in the field. Quadrats 2 and 11 are located
to the south of a four metre high hawthorn hedge (2) and along a shrub of
creeping willow (11). The remaining quadrats lie distributed over the terrain,
among vegetations of types II, III, and IV. Along the edges of the hedge and the
shrub there is a rather well-developed strip of grass in which older nymphs and
adults were observed throughout the summer, the vegetation apparently being
extremely suitable for habitation.
Regarding the situation in the vegetations of type I, we may state that M. macu-
latus hatches in it in large numbers; that the numbers of nymphs of the later
immature stages are low when this vegetation is surrounded by types II, III, and IV
but that nymphs and adults can maintain themselves if there is a protective
vegetation in the immediate vicinity; and that while the adults are initially scarce
they nevertheless occur in larger numbers later in the season.
Type II. The vegetations grouped under type II are distinguished from those of
type I by the development of the moss layer. The high herb layer gives slightly
more coverage than that of type I. The results of the counts on Quadrat 7 are
given in Table 21.
Here, too, as in the type I quadrats, many nymphs are hatched. However, the
numbers of specimens of later instars found later in the season are appreciably
smaller. Adults were scarce until late in September. On north slopes, on which
there are about 150 sq.m of uniform vegetation, very few grasshoppers were
observed during the three years of the investigation.
Type III. The vegetations belonging to type III are characterized by a dense
moss and lichen layer and grass tussocks in the high herb layer. This type was not
sampled in 1955. The results of the counts are given in Table 21.
Nymphs hatched on all quadrats. Hatching was observed on small, bare spots
under blades of Festuca grass and in places where ants regularly brought sand to
the surface. Because a large number of quadrats in this type could not be sampled
on 14 May, 1954, little can be said concerning the numbers of the first instar
hatched and later present on them. On 13 May, 1954, however, in connection with
the study of the hatching sites of M. maculatus, three quadrats measuring one
sq.m in type III vegetation were sampled (EE, FF, and CC, Table 11). On these
quadrats, 13, 65, and 49 first instar nymphs were found, respectively. The quad-
rats measuring one sq.m were located in the same part of the terrain as Quadrats
46, 47, 48, and 50. These data, taken together with the large numbers of second
instar nymphs found on the quadrats, justify the assumption that the numbers
of nymphs hatched on the unsampled plots were large.
Nymphs of the third and fourth instar remain in type III vegetation in larger
numbers than in types I and II. Adults are found primarily in the summer months.
During the sampling of these quadrats in July and August on warm, sunny days,
most of the older nymphs and adults were seen in or close to the grass tussocks.
Summarizing, we may state that many nymphs hatch in type III vegetation but
also that the sharp decrease in numbers that occurs in types I and II is not seen
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
404
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406 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
to the same extent in type III. Type III vegetation evidently induces a population
of more stable size.
Type IV. The vegetations grouped under type IV possess a moss layer covered
by a rosette layer and a dense carpetting layer. The results of the counts in this
type of vegetation are given in Table 22.
M. maculatus was consistently observed on all the sampled quadrats during a
large part of the season. Nymphs were hatched although not in such high numbers
as were reached on types I and II, the typical hatching sites. Older stages remain
present in relatively large numbers, while the adults populate the quadrats in
varying density.
Type V. The vegetations included in type V are characterized by the large
proportion of grasses in the high herb layer. The results of the counts are given
in Table 23.
The numbers of nymphs hatched in this type of vegetation are very small, and
the first instar is later poorly represented as well. Quadrat D formed an exception
in this respect in 1955. In 1954 the situation in this locality was quite different
from 1955, which is important for the evaluation.
In the months June and July the quadrats were populated by large numbers of
second, third, and fourth instar nymphs and adults. Since repeated observation of
these vegetations showed that few or no nymphs hatched, the increase in the
numbers of older nymphs and adults must be ascribed to migration from less
dense vegetations.
The adults, which are initially numerous in this type of vegetation, either occur
rarely or are entirely absent from the middle of August on. This phenomenon was
particularly distinct in 1954, in a period in which appreciable numbers of nymphs
still populated the field.
The type IV quadrats were chosen in the terrain such that they were located not
only in the transitional areas between high, dry parts and lower, damp parts but
also in the three to four metre wide strips along the hedges surrounding the ter-
rain. In one part of the terrain along a hawthorn hedge in which Quadrats 3, 4,
and D were chosen five, three and one metres respectively from the edge of this
vegetation, large numbers of nymphs and adults occurred in the summers of 1953
and 1954. In the summer of 1955, due to the early and vigorous growth of the
grasses, the vegetation of this part of the terrain was so changed in structure that
a large part of the cover outside the one metre wide borders had to be assigned
to type VI. The adults were found in this year almost exclusively in these borders.
In 1956 both the vegetation and the M. maculatus population in this part of the
terrain were comparable to those in 1954.
A similar border population” was observed outside the terrain of investigation
in a type V vegetation occurring in a corner of the landward dunes about 15 metres
in width. Older nymphs and adults of M. maculatus were found in 1954 in the
approximately five metre wide border, although few or no other grasshoppers were
found in this vegetation.
407
B. M. LENSINK : Distributional ecology of Acrididae
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408 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Type VI. Type VI vegetation includes the high-growing, dense grass and herb-
rich dune meadows. Few or no M. maculatus were found in the eight quadrats
chosen in this type of vegetation. Three quadrats, however, lay in this vegetation,
which became type III with almost no transition.
The situation on these quadrats is comparable to that in type V: no hatching;
migration of older instars from outside the quadrats. It was noted for one of the
quadrats, located in the eastern corner of the terrain, that after a period with
strong west wind more adults were observed in the border zone than elsewhere,
a case for which the direct influence of macroclimatological conditions may be
assumed to the exclusion of microclimatological influences.
Summary. Summarizing the results of the observations during the period in
which M. maculatus populated the field, we may state the following:
(a) The greatest numbers of nymphs hatch in vegetation types I and Il; in
places where the moss layer is only partially developed (type I), unusually high
numbers hatch;
(b) Nymphs hatch in types III and IV vegetation, but in smaller numbers
than in types I and II;
(c) Nymphs hatch in types V and VI only in very small numbers;
(d) In types I and II the numbers of nymphs drop rapidly a month after
hatching; few nymphs of the third and fourth instar occur; adults are scarce,
although there is reason to assume that they are relatively more numerous later
in the season;
(e) In types III and IV the numbers of nymphs decrease during the early
summer, but at an appreciably slower rate than in the types I and II;
(f) During the first month after the beginning of hatching, nymphs of the
second and older instars enter type V vegetation, and this also occurs in the bor-
ders of vegetations of type VI;
(g) Later in the season, in about the middle of August, no adults are observed
in vegetation of type V and in the borders of type VI.
Results. The study of the occurrence of M. maculatus has shown that the occur-
rence of grasshoppers cannot be determined by means of one or more incidental
observations. We have seen that in the course of the season in which this species
populates the field, the places in which the most nymphs hatch become empty,
while older nymphs appear in the dense, herb-rich vegetations. Migration thus
occurs in the course of the season from less dense to denser vegetations, evidence
of a shift in the choice of the habitat.
The results of this migration from types I and II to types III and IV is hardly
or not at all noticeable in large parts of the field because the surface of the terrain
covered by type I and II vegetation is very small in comparison to the surface
covered with types III and IV, and the spread is consequently very great. Only
in places where there are many grasshoppers, i.e., where vegetation types I and II
lie close to types IV and V, can this shift be clearly demonstrated, as is the case,
for example for Quadrats 2, B, 3, 4, and D.
Later in the season, when mostly only adults are present, the insects disappear
409
B. M. LENSINK : Distributional ecology of Acrididae
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410 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Myrmeleotettix maculatus
type I 1954
quadrat 9
type I 1954
quadrat 7
type I 1954
quadrat 47
type U 1954
quadrat 21
| 1 2
instar BO
4
7 12
8 28
sept. oct.
Bane NA AT NES Tee 20 29! 6
Graph 1. Myrmeleotettix maculatus. Abundance of the various stages on the sampling dates.
Absciss: dates on which the samples were taken; ordinate: number of specimens. Further
explanation in text. Continued on next page
B. M. LENSINK: Distributional ecology of Acrididae 411
Myrmeleotettix maculatus
oe type x 1954
20 quadrat 3
10 : |
DAG le DE Ln PLE DE) STE TESTEN MANE
i i AOS
april may june july august Sept oet
Graph. 1. Continued from p. 410
from the dense, herb-rich vegetations. An increase in the numbers of adults in the
less dense vegetations is difficult to demonstrate, because in this period the density
of the insects has already passed its maximum and the spread is again very great.
It was nevertheless striking that, for instance, in Quadrat 9, there was a distinct
increase in the number of adults. This Quadrat was located at a distance of about
30 metres of type V vegetation. For purposes of illustration, Table 24 gives the
numbers of adults found on type I and type V quadrats in 1953, 1954, and 1955.
Graph 1 shows the variations in population in 1954 in a number of quadrats
belonging to types I—V, also for illustrative purposes.
The distribution of C. biguttulus
As has already been said nymphs of C. biguttulus were found to hatch in
vegetation types I—V. The majority, however, hatched in the very thin vegetations
of type I, particularly where this type lay in the vicinity of types V and VII.
The numbers of insects of this species observed in the terrain were consistently
small. In 1955 the numbers were so small that conclusions can barely be drawn.
The largest numbers (63 nymphs) were found on 5 July, 1955 (on 23 quadrats
measuring three sq.m each).
If, as for M. maculatus, we compare the total numbers of nymphs of each of
the four instars and the adults found on the quadrats of each type of vegetation
during the season in 1954, a general picture of the variation in numbers is again
apparent. The figures are given in Table 25.
Table 25. The total numbers of nymphs and adults (a) of C. biguttulus found on the
quadrats of each type of vegetation during the entire season in 1954.
Instar 1 2 3 4 A No. of quadrats
sampled
Type I 48 28 3 2 4 5
Type I 79 38 8 2 11 2
Type II 107 5 2 1 1 3
Type III 119 91 16 6 11 8
Type IV 105 97 35 20 35 9
Type V 34 48 59 40 39 8
Type VI 18 43 57 29 31 11
No. of samplings 10 10 9 9 10
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
412
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B. M. LENSINK : Distributional ecology of Acrididae 413
In this table a correction has been applied for the first instar on the five
type I quadrats, because of the partial loss of 14 May, 1954 sampling (p. 398).
The numbers of the first instar missed on type I vegetation are estimated to be 1/3
of the numbers observed during the other samplings.
Quadrats 2 and 11 were sampled on 14 May and the data from these quadrats
are given separately in the table because by far the largest numbers were found on
these two quadrats. The numbers of the first to fourth instar decrease strongly
in vegetation types I and II, the average figures per quadrat in type I being 1.81,
0.94, 0.17, 0.06, and in type II 0.56, 0.17, 0.08, and 0.04. These figures clearly
reflect the limited density.
In the type III tussock vegetation and the type IV herb-rich vegetation, the
numbers also drop during development, but not with the initial rapidity found
for the two other types. For type III the average figures are 1.50, 1.14, 0.22, and
0.08 and for type IV 1.16, 1.08, 0.43, and 0.25 nymphs per quadrat respectively.
In the dense, low herb-rich vegetation of type V and the dense, high grass cover
of type VI, very few nymphs of the first instar were found initially, but the
numbers increased with the more advanced instars. Per quadrat the average figures
for type V are 0.42, 0.60, 0.82, and 0.58 and for type VI 0.16, 0.40, 0.58, and
0.29 nymphs respectively.
Comparison of these values indicates that in the types I and II vegetations the
numbers of nymphs drop more rapidly during development than in types III and
IV and that there is an initial rise in types V and VI, followed by a drop.
As for M. maculatus, we may state that in addition to mortality there will be
an effect due to migration from the microclimate of vegetation types I and II
(where conditions tend to become extreme in the course of the early summer)
to vegetations that offer more protection, i.e, types III through VI. The fact that
migration is not noticeable in types III and VI may be explained on the basis of
the fact that both the latter types of vegetation constitute an appreciably larger
proportion of the terrain than the former. The spread is consequently very great.
The data for the individual quadrats will not be discussed because the numbers
are so low, especially those for the older stages, but the general tendencies in the
individual types will be evaluated.
Types I and II. Table 26 gives the total numbers of nymphs of all instars and
the adults found on the 1954 sampling dates on seven quadrats of type I and three
quadrats of type II. Table 29 gives the same data for seven type I quadrats in
1955. These tables show that the quadrats with a very thin cover are populated
principally by nymphs of the first and second instar, the older instars being found
only in small numbers. Beginning in July, this species has almost disappeared
from these quadrats. Adults are not seen on them until the end of August, al-
though in 1954 they were seen on 29 June and in 1955 on 11 July, during field
sampling.
Quadrats 2 and 11 showed the largest numbers in 1954 in comparison with the
other quadrats of the same type. The location of these quadrats is exceptional, next
to a hawthorn hedge and creeping willow shrub (see for M. maculatus p. 403)
in which and beside which adults of C. biguttulus wete seen throughout the sum-
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
414
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B. M. LENSINK : Distributional ecology of Acrididae 415
mer. This species also showed the highest numbers on Quadrat 2 in 1955. The
vegetation along the edge of low shrubs apparently provides a very favourable
habitat for this species.
Types III and IV. Tables 27 and 30 give the results of the counts in 1954 and
1955 in the same way as the tables for types I and II. Type III was not studied
in 1955. First and second instar nymphs predominated on the quadrats of both
types. Here again the numbers of the third and fourth instar nymphs are distinctly
lower. Adults appeared first at the beginning of August in 1954 and the end of
August in 1955.
Types V and VI. Tables 28 and 31 give the results of the counts in 1954 and
1955. First instar nymphs were observed on these quadrats too. In comparison
with the figures for the other types of vegetation, the numbers in type VI are
appreciably lower.
Experiments concerning the choice of oviposition substrate have shown that the
uninterrupted vegetation and the insulating layer of dead grass are not suitable
for oviposition (LENSINK, 1962). In the high, dense grass vegetation an egg pod
of C. biguttulus was occasionally found in old molehills (which occur here and
there in low- -lying spots), and this explains the observation of newly-hatched
larvae of this species in this type of vegetation.
In 1954 nymphs or adults were found throughout the whole season in both
the low dense (V) and the high dense (VI) grass vegetations. The numbers of
the second, third and fourth instar are higher than that of the first instar, in-
dicating migration from the less dense vegetation in which the larger numbers
of the first instar are found. This is also apparent from the totals of all the qua-
drats per type.
This migration is hardly noticeable for 1955 in the tables. It should be remarked
here that in that year the grass cover of types V and VI showed a rapid growth
early in the summer in comparison with 1953 and 1954, so that the structure was
unusually dense, particularly in type VI. Migration reached only the edges of these
vegetations. The same phenomenon was observed in Quadrats D and 4, located
in the same area along the hawthorn hedge as Quadrat 2.
No reduction in the numbers of insects in these dense types of vegetation, such
as that found for M. maculatus in type V, was seen for this species. Fewer speci-
mens were seen and heard towards the end of August in the edges of low creeping
willow shrubs. This could not be demonstrated by sampling because the densities
had already become very low. These observations point to a shift to more open
vegetation, which is also expressed in the data for the quadrats of the first four
types of vegetation at the end of the season and the large numbers of newly-
hatched larvae in this types in spring.
A comparison of the numbers of adults found on the various dates of sampling
in the quadrats of the various vegetation types fail to explain this shift conclusively.
Summary. Ia the period during which C. biguttulus occurs in the terrain in
which the investigation was carried out, we see the following:
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
416
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B. M. LENSINK : Distributional ecology of Acrididae 417
(a) The insects hatch in all types of vegetation, but the largest numbers hatch
in type I where it borders on type V or type VI or along the hedges locally bor-
dering the terrain and beside growths of creeping willow (VII);
(b) During development the numbers of nymphs in types I and II decrease
and few or no specimens of older instars occur in these types during the summer;
a decrease is obvious in types III and IV, although during the summer
older instars do occur, albeit in limited numbers; in types V and VI in 1954 an
increase in the numbers of the second and third instar was observed in relation
to the numbers of nymphs hatched in this type of vegetation; in 1955 this increase
was not observed in type VI (probably as a result of the very rapid growth of
the vegetation which became dense very early in the season) ;
(c) Adults were not found in the quadrats of types I, II, HI, and IV until late
in the season (at the end of August) in 1954 and 1955; in the remaining quadrats
in both dense vegetation structures they were found from their first appearance
in the field until the period of the night frosts in October.
Results. The investigation into the distribution of C. biguttulus again showed
that incidental observations are insufficient to establish the occurrence of the
species from egg to adult. And shifts occur for this species also. The oviposition
sites are located in entirely different parts of the field than are the places in which
the adults live.
There is distinct migration towards the dense, herb-rich growths and grass
vegetations. The numbers of adults are limited and they are found primarily in the
denser vegetations. Only later in the season do they occur elsewhere, in the types
I—IV vegetations. These latter insects could not have developed from a locally
present population, so they must have shifted from the more dense to the less
dense vegetation.
The distribution of C. albomarginatus
The investigation into the hatching sites of C. albomarginatus showed that the
first instar was present in all the vegetations of the terrain. Since the nymphs are
not very mobile during the days immediately following hatching, it may be
assumed that they had hatched where they were found and that they later spread
out from these hatching sites. The largest numbers of older nymphs were found
in the denser grass vegetations, in 1953 and 1954 especially in types V and VI
and in 1955 especially in type IV.
Tables 32 and 33 give the figures for the nymphs and adults in 1954 and 1955.
The "14 May, 1954” correction (p. 398) was applied to Table 32. The numbers
of nymphs of the first instar in types V and VI were augmented by 50 on the
basis of the data obtained from the 22 quadrats which could be sampled on
14 May.
In both 1954 and 1955 there were few nymphs and adults of this species on the
very thinly-covered places (types I and II). In 1954 the numbers were much larger
in type III, although they rapidly decreased. The figures for the first to the fourth
instar were 1.52, 1.23, 0.08, and 0.02 nymphs per quadrat respectively.
418 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Table 32. The total numbers of nymphs and adults (a) of C. albomarginatus found on the
quadrats of each type of vegetation during the entire season in 1954.
TOStar 1 2 3 4 A No. of quadrats
sampled
Type I 11 2 2 0 0 7
Type II 3 3 0 0 0 3
Type III 73 69 6 2 4 8
Type IV 183 81 21 10 51 8
Type V 100 86 33 26 25 9
Type VI 233 509 489 475 436 11
No. of samplings 6 7 9 10 12
Table 33. Same observations as Table 32, but in 1955.
Tnstar 1 2 3 4 a No. of quadrats
sampled
Type I SJ 35 13 5 3 7
Type II 10 3 0 0 1 1
Type III — — — — _- —
Type IV 280 210 46 17 61 6
Type V 8 24 12 14 11 3
Type VI 7 45 52 84 118 6
No. of samplings 7 7 7 4 8
Larger numbers were initially found for the first instar in type IV. In 1954 the
numbers of the older nymphs decreased rapidly, i.e., from the first to the fourth
instar the averages were 3.81, 1.45, 0.29, and 0.13 nymphs per quadrat respectively.
In 1955 the decrease took place only after the second instar. This undoubtedly
related to the prolonged cold spring. The average figures were 6.67, 5.00, 1.09,
and 0.71, respectively. More adults occur in this type of vegetation than in the
three previously mentioned types.
No large numbers of this species were found in type V in 1953, 1954, and 1955.
The figures for the first to fourth instar in 1954 were 1.85, 1.36, 0.48, and 0.29
and in 1955: 0.38, 1.14, 0.57, and 1.16. In the terrain this vegetation type is
found beside hedges and low shrub, surrounded by a thin grass cover and con-
stitutes only a small proportion of the terrain.
The numbers of nymphs found in type VI show the most interesting variations:
the average figures in 1954 were 3.54, 6.60, 4.94, and 4.74 nymphs per quadrat
respectively, and in 1955: 0.17, 1.07, 1.24, and 3.50. In both years a distinct
increase in the numbers of the second instar was observed, caused by migration
from less dense vegetation. The numbers of the older instars are much larger than
in types I—V.
Types I and II. Very few nymphs and adults of this grasshopper species occur
in the vegetations with a broken or unbroken moss cover and a very thin grass and
herb cover. The numbers of insects on the quadrats with these growths are given
419
B. M. LENSINK : Distributional ecology of Acrididae
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B. M. LENSINK : Distributional ecology of Acrididae 421
in Tables 34 and 39, from which it can be seen that neither type provides a suitable
habitat. Only a small number of nymphs hatch there and the numbers of older
instars and adults are small to very small.
More of the second and third instar occurred in 1955 than in 1954. If it may be
assumed that the developing nymphs in these vegetations migrate in response to
extreme microclimatological conditions occurring in the early summer and the
absence of adequate shelter, the prolongation of this migration over a longer
period in 1955 may be attributed to the relatively low temperatures in the early
spring and early summer of that year, as a result of which extremely high tem-
peratures did not occur until later in the year.
In 1955 adults first were found after the middle of August. In 1953 and 1954
the adults found on all of these quadrats were also first seen only after the middle
of August. If migration occurred from the denser to the less dense vegetation, it
would explain these small numbers. For the total number of adult insects in the
field is in the process of decreasing strongly in the middle of August, while almost
all the quadrats of both types are located at a distance of 50 metres from the
growths in which the most adults live during the summer.
Type III. In the quadrats covered with vegetations in which tussocks of Festuca
ovina dominate heavily, the distance separating them from a high, dense grass
cover (type VI) plays an important part in determining how many of this species
are present. Table 37 gives the results of the 1954 counts (no inventory was made
of this type of vegetation in 1955). Quadrats 39 and 44 lie at a distance of five to
ten metres from the edge of the dense vegetations, and the other quadrats are
separated from them by more than twenty metres. The numbers are not very large,
but the difference is obvious.
These vegetations are also hatching sites of nymphs which are rarely or never
seen in them from the beginning of the third instar. Adults are also almost never
seen. In 1953 they were seen in small numbers much later in the season; in 1954
and 1955 the figures were negligible.
Type IV. Nymphs of C. albomarginatus occur locally in this type of vegetation.
Tables 38 and 40 give the data for 1954 and 1955, respectively. The quadrats
with the most nymphs, including especially the first, second and third instars, lie
in the vicinity of vegetations belonging to type VI in which especially adults are
found in the summer (1954, Quadrat 5 and 1955, Quadrats 5, H, and J, in Tables
38 and 40, respectively). The other quadrats of this type are located more than
20 metres away of the edge of a high, dense grass cover. Only very small numbers
of nymphs were seen on these quadrats.
In 1953 sampling was not begun before the second half of June, a time at which
in all three years few nymphs were still present. The fourth instar was seen in
small numbers; in 1953 and 1954 adults were seen during August on the sampled
quadrats of this type. This is remarkable because the first adults were seen in the
field as early as 20 June in 1953 and on 14 June in 1954. In 1955 the first adults
were not seen in the field until 5 July, while they were found in type IV on 25
July during the sampling. Their late appearance in 1955 is attributable to the
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
422
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B. M. LENSINK : Distributional ecology of Acrididae 423
retarded development of the nymphs caused by the low temperatures during the
spring.
Since it is excluded (certainly in 1953 and 1954) that the adults developed from
the first instars present on these quadrats in the spring, this must have been a case
of migration from other, in this case denser, vegetation. The low numbers are
explained by the fact that here, too, the population throughout the terrain is
becoming reduced. In addition, the surface of the terrain covered with type VI
vegetation is appreciably smaller than that covered with less dense vegetation
types. In other words, the spread is very large here, so that the chances of finding
adults later in the season in type IV, on the small numbers of quadrats are very
small.
Type V. In this vegetation, very few nymphal stages and adults were seen in
1953 and 1955; in 1954 rather more were observed. This can be clearly seen from
Tables 35 and 41, which give the numbers found on the various quadrats belong-
ing to this type, in 1954 and 1955.
The adults were seen primarily in the summer months of July and August.
Later, in September, they appear only sporadically. In view of the findings in the
less dense vegetations, we may assume that the adults migrate out of type V
vegetation during the month of August.
Type VI. During a large part of the season C. albomarginatus is found (some-
times in large numbers) in the high, dense grass cover of this type. This vegetation
occurs in the lower, damp parts of the terrain, especially along hedges. In the
lower parts the height of the grass cover is rather even, and large tussocks of grass
alternate with a lower herb-rich cover. On the higher parts Calamagrostis epigejos
dominates, forming an unbroken cover. At first, in winter and early spring, the
grasses that form by far the most important part of this type of vegetation still lie
flat on the thick layer of dead grass which covers the soil. During the spring,
sparsely scattered blades of grass begin to shoot up. In May and June, however,
when the grass is 30 to 40 cm above the dead layer, it forms a complete cover.
In the time during which it flowers, and the flower stalks stick out above the
cover, the vegetation often does not become higher because the wind tends to
flatten it. This creates a thick mat of grass on the ground. In the lower parts of
the terrain, herbs flower locally between the grass clumps.
Tables 36 and 42 give the numbers of nymphs and adults, as found from the
various data of 1954 and 1955. Differences between 1953 and 1954 on the one
hand and 1955 on the other, require separate discussion.
In 1954, newly-hatched larvae were found on all the plots. Except for Quadrat 6,
their numbers were consistently very small. The smaller nymphs can still be found
in the as yet rather open grass cover in May, although extreme care had to be used
in counting a three sq.m quadrat. It seems to me unlikely that the small numbers
of the first instar are to be attributed to errors in the sampling technique.
The numbers of the second and third instar are higher for almost all quadrats
than the numbers of the first instar. Because mortality during development reduces
the numbers of nymphs, this increase must be seen as the result of the arrival of
424 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
nymphs from less dense vegetations to these high, dense growths of grass. The
adults occur in these vegetations from the beginning of July on.
As for the type V quadrats (but here to a greater extent) the numbers drop
sharply after the middle of August. Here, too, migration to the surrounding thin
vegetations must be considered.
In 1953, before the start of sampling, numerous nymphs of this species were
found in the high, dense grass cover. In this year, too, the adults were pre-
ponderantly found in this vegetation. After the end of August, however, the
numbers of adults dropped sharply while an increase was found in other places.
This phenomenon was clearly seen again in the part of the terrain in which Qua-
drats 6, F, G (type VI), and 5, H, and J (type IV) (Table 40) were located.
During the years of the investigation, this area had very large numbers of this
species. The high, dense grass cover is dominated by Calamagrostis epigejos which
grew very high (70 cm). Quadrats 6, F, and G lay about one metre within the
border of this vegetation. Quadrats 5, H, and J were sampled over two to three
metres from the outer border of the above-mentioned vegetation, being themselves
covered with a type IV vegetation. Graph 2 gives an illustration of the situation
on these quadrats.
As early as 1953 it was evident that relatively large numbers of adults of C.
albomarginatus occurred in Quadrat 6. The numbers dropped in the second half
of August, while there was a distinct increase on Quadrat 5 in this period. In 1954,
nymphs hatched in Quadrats 5 and 6. In 6, the numbers of older instars increased,
but in 5 it dropped markedly. The numbers of nymphs were sharply reduced in 6
during August, but in 5 they increased sharply. In 1955 the numbers of quadrats
were increased in order to collect more data on this phenomenon. In that year
few or no nymphs of the first instar were observed in Quadrats 6, F, and G,
although eggs had been observed during the winter. In all probability, this
phenomenon may be attributed to the effect of the very low night temperatures in
the spring of 1955. In comparison with 1954, the numbers of older nymphs
Chorthippus albomarginatus
type U 1953
10 quadrat 5
type VI 1953
quadrat 6
01.3204
instar B O @ B B
ea MZ Bl Oe) ol 8
nes 1 Eken 200, 3721 oT eS SH 7 14 24 2 7 IS 27
June july august sept. oct.
Graph 2. Chorthippus albomarginatus. Abundance of the various stages on the sampling dates.
Absciss: dates on which the samples were taken; ordinate: number of specimens. Further
explanation in text. Continued on next pages
B. M. LENSINK : Distributional ecology of Acrididae 425
100 Chorthippus albomarginatus
type IX 1954
quadrat 5
100
90
80
70
60
50 9 gs type VI 1954
7 di quadrat 6
40 4 7 1 2 3 4 adult
g (È instar B_D 8
2 gi
AG È
BE 3
20 i Hi
GE di
Ad 9%
gs ZE
10 Zi GE
a À ii |
PARI AA 29 CHAMPS NS
april may june august Sepimocti
Graph 2. Continued from p. 424
remained small, and development was greatly retarded. The numbers of adults
were not large, and they disappeared almost entirely from the sampled quadrats
during the course of August. In Quadrats 5, H, and J of type IV, nymphs hatched.
Adults were found there rather late in the season, although they were first
observed on 5 July, 1955 in the dense, high grass cover.
426 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Chorthippus albomarginatus
type W 1955
quadrat 5
type Iv 1955
quadrat J
10
O
30 type m 1955
quadrat H
type VI 1955
quadrat 6
10
(o)
20 type VI 1955
quadrat F
type VI 1955
quadrat G
; ı 2 sì a eis
instar B O 8
FEINEN Ones agio mt en Vs Na. 2%
april may june july august sept. oct.
Graph 2. Continued from p. 425
We may conclude from these data that eggs of C. albomarginatus are laid in
both the dense grass cover and in the thinner vegetations. In spring, the second
and third instar nymphs migrate to the denser and more protective vegetations,
where the largest numbers are concentrated during the summer and the females
B. M. LENSINK : Distributtonal ecology of Acrididae 427
lay their eggs. Later in the season, in the second half of August and later, the
adults migrate because the microclimatological conditions begin to deterioriate in
the dense grass cover, shifting to the surrounding vegetations which are less dense
and relatively warmer, in which they also oviposit.
The same phenomenon is seen in 1955, but the picture is somewhat blurred
because as a result of the lower temperatures in the spring and early summer,
migration was apparently less pronounced. The rapid growth of the grass, espec-
ially in type VI vegetation, also contracted the transition zone between the types
of vegetation. In the summer of 1955, the largest numbers of adults were also
found in this border zone.
Summary. If C. albomarginatus is followed during the season in this terrain, the
following is observed:
(a) As soon as the eggs of this species hatch, the first instar nymphs are found
in almost all types of vegetation;
(b) Very small numbers of nymphs hatch in vegetation types I and II; few
are found in types III and IV except were they border on a high, dense grass cover
in which many of the grasshoppers are seen during the summer; most of the
nymphs hatch in types V and VI;
(c) In types I—IV, the numbers of nymphs are rapidly reduced during develop-
ment, and few or no adults are seen during most of the summer;
(d) In types V and VI the numbers of the second and the third instar increase;
in July and August the adults populate these types of vegetation;
(e) Later in the season (in general, starting in the middle of August) very
small numbers of adults appear in types I, II and III, and somewhat larger num-
bers in type IV, while in the same period in types V and VI the numbers of adults
drop or they disappear entirely.
Results. Investigation of the occurrence of C. albomarginatus in the vatious
types of vegetation distinguished here shows that this species and its stages
occur not only in the high, dense, relatively moist grass cover but also in the drier,
thinner vegetation. The hatching of the larvae was observed in many types of
vegetation.
During its development, this species was concentrated in the denser growths.
The denser grass vegetations show an increase of the older instars. Adults are first
observed late in the season in types I—IV, although they have long been present
in the dense types of vegetation. There is evidently migration from the dense back
to the less dense vegetation.
THE ACTIVITIES OF THE GRASSHOPPERS DURING THE DAY
Introduction
The preceding chapter has dealt with the activities of the various nymphal
instars and the adults of the three species of grasshoppers during the season. The
present chapter is concerned with the activities of the grasshoppers during the day.
428 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
Attention was drawn to the daytime activities of these insects because, especially
for M. maculatus, it was observed that at particular places (Quadrats 2 and 11)
there was a diurnal variation in the density of the population. What role is played
here by the presence of a hedge or a shrub? Do the insects make use of such cover?
We were unable to examine such factors for C. albomarginatus and C. bigut-
tulus, but a description of the way in which the insects live in the vegetation will
serve to clarify the influence of a number of factors that affect the choice of the
area in which to live.
The investigation of the activity of the grasshoppers during the day was carried
out primarily in 1955. Observations were carried out in various types of vegetation
under different weather conditions, sometimes several hours in duration. Attention
was given to such behaviour as: eating, sunning, walking, jumping, courting,
mating, egg-laying, and reactions to enemies. Special attention was given to the
place and height at which the insects were found in the vegetation. All these
activities are strongly dependent on the state of the weather. During rain or cold,
cloudy weather the insects hardly move at all. With sunny, warm weather they are
often extremely active, and are easily disturbed so that observation must be done
unobtrusively. Observation was done from places where a number of grasshoppers
could be seen simultaneously. The data collected for the three species will be dis-
cussed successively. The observations concern primarily M. maculatus and C. albo-
marginatus. C. biguttulus is extremely difficult to follow because especially the
older instars and the adults are extremely active and sometimes range over large
distances.
The diurnal activities of M. maculatus
During the morning this species shows almost no activity, in either the thin or
denser vegetations, in the spring (May, June) and fall before 9:00 a.m. The
nymphs and adults sit quietly on the ground or between the leaves of the herbs.
As the day becomes warmer and the solar radiation begins to penetrate the
vegetation, the grasshoppers begin to show activity; they walk and eat and the
males begin to stridulate. As the morning advances, these activities increase. The
males pursue the females, and courting becomes intense. The nymphs and adults
both seek favourable places from which to profit from the sunshine. Oviposition
is observed only in places where favourable substrates (especially type I) border
on a more protective vegetation, and these are also the most heavily populated
parts of the field. During the afternoon, as the sun reaches a lower position and
the angle of its rays reduces their penetration, the activity subsides. In the spring
and fall, activity ceases between 4:00 and 5:00 p.m. and during the summer
months between 6:00 and 7:00 p.m. Only once have I heard a soft singing in a
type IV vegetation around 9:30 p.m., on an occasion of high temperature and
little wind.
Thus on warm sunny days there is high activity throughout the day. On days
with changeable weather, activity increases rapidly during the sunny periods, to
drop as rapidly when a cloud passes over the sun. These changes are accompanied
by fluctuations in the intensity of the song. It is of interest to mention here that
B. M. LENSINK : Distributional ecology of Acrididae 429
during the eclipse of the sun in 1955 a distinct decrease in activity was observed.
On dark days activity is very low. With a cloudy sky and temperatures around
17° C (measured in the weather shed at an altitude of two metres) only scattered
soft song was heard and only a few eating insects observed. In thís connection it
may be mentioned that the activity of the grasshoppers held in cages in the
laboratory showed a distinct increase when a lighted electric bulb was hung in
the cage; singing and courting followed directly.
Because a sufficiently large population occurs throughout the season in vege-
tation types III, IV, and V, the diurnal observations were done systematically in
them. With favourable weather conditions, more than 40 hours of observation were
done in these types during various times in the season. Incidental observing was
done during other weather conditions. The most active insects are the males. The
nymphs and the females show little activity. They walk short distances, now in a
sunny place and then again in the shadow of a grass tussock or the leaf of one
of the many herbs. During the middle of the day, when the solar radiation
penetrates deep into the vegetation, however, it is observed that many nymphs
and adults are found not on the ground but some distance above it on a blade of
gtass or a leaf.
A large number of grasshoppers were observed for two minutes per insect.
The place in which they were located in the vegetation under various weather
conditions was noted. These observations clearly showed that with strong radiation,
during the middle of the day, most of the animals were found at a height of one
cm or more in the vegetation. With alternating sun and cloudiness and/or much
wind, most of the insects were seen on the ground. Before 10:00 a.m. and after
4:00 p.m., the insects predominantly sat on the ground sunning themselves. These
data, with the locally-measured temperatures, are given in Table 43.
Table 43.
| Temperature No. of insects
Type of weather
| 1 cm above ground | on ground 1 cm high or higher
strong sunshine ZONE 18 56
sunshine/cloudiness/wind 26—30° C 23 3
before 10 a.m. & after 4 p.m. 25€ 247, 8
With bright sunlight, the temperature close to the ground is appreciably higher
than a few cm above it. The behaviour observed proves the fact that these animals
also show a vertical migration. The assumption that by this migration they
attempt to avoid extreme temperature, is very probably correct. In the tussock
vegetations (type III), in sunny weather, we found the most grasshoppers during
sampling in the tussocks on the grass blades at about five cm above the ground.
Here again the insects apparently avoid the high temperatures above the moss
layer and choose milder places where the temperature lies only a few degrees
higher than the air temperature above the grass. For purposes of illustration an
excerpt of the field-notes of 24 June, 1955, may be included here; it serves as an
example of the behaviour in type IV vegetation in sunny weather with a weak to
430 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
moderate N-NW wind and occasional high-altitude clouds. The observation time
was between 11:10 and 11:45 a.m. A male M. maculatus was followed:
11:10 a.m. — with flank partially in the sun on the rosette leaf of Hypochaeris
radicata — shifts with the head towards the sun / 11:15 — entirely in sun on
leaf, remains immobile till 11:21 — turns with head towards sun, creeps into
shadow of leaf of Calamagrostis epigejos, again turns half of flank into sunlight /
11:26 — walks ten cm towards shadow of tussock of Festuca ovina / 11:29 —
walks, jumps fifteen cm further along the ground, sits half in shadow with head
in direction of sun, against a leaf, turns flank towards sun / 11:32 — stridulates
in this position, walks further, eats on shady side of Festuca ovina (sun clouded
over), insect leaves shadow (sun returns, veiled), insect turns with flank in sun /
11:40 — (sun gone) head in sun, (sun back) sun on flank / 11:44 — creeps in
shadow of clump of Festica ovina...... 11:45.
Similar observations were consistently made for nymphs and adults in vegetation
types III, IV, and V. The most striking point is that they alternately sit in the sun
and then in the shade of a leaf or stalk. No such opportunity is offered by the
more open vegetations (types I and II), in which the environment is extreme and
which are therefore almost unpopulated during the summer months.
In places where types I and II vegetations border on type V or VII (the most
populated parts of the terrain), the behaviour is even more remarkable. In these
very thin vegetations, in contrast to other parts of the field, nymphs and adults
are seen regularly. A series of continuous observations were made in this kind
of vegetation. One of the observation points chosen was a small terrain with a
low Salix repens growth along a sttip approximately one metre wide, of almost
bare sandy soil. On the other shide of this strip there was a vegetation of type IV
with transtition to type V. M. maculatus and a few C. biguttulus were pre-
ponderantly observed here, with an occasional specimen of O. coerulescens.
The observations were made in a plot two metres long and one metre wide of
almost bare sandy soil for a total of six days, representing 58 hours (Tables 44
and 45), by two observers in a position to note all grasshoppers going or
coming between types VII and IV, and the stretch of sand. The total
observation time was divided into half-hour periods. For each period notation was
made of the numbers of insects present on the strip as determined by the numbers
which left or entered the strip, as well as their activities such as eating, oviposition,
walking, etc. Notes were also made of the behaviour of the grasshoppers present
in the Salix growth. Table 44 gives the numbers of males and females observed
in the strip in the successive half-hour periods, the numbers of females which dug
into the soil with their abdomens while searching for oviposition substrates, and
the numbers of females which then laid eggs. The cloudiness in the various periods
is indicated by the letters A—D.
The weather conditions on the different days were:
25 July — moderate NW wind, sunny, occasional high-altitude clouds, in the
afternoon increasing cloudiness, after 6:30 p.m. very cloudy with some rain;
26 July — moderate NE wind, half overcast;
27 July — sunny, warm weather, clear, later in the day some cloudiness;
431
Distributional ecology of Acrididae
B. M. LENSINK :
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432 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
28 July — rather strong N-NE wind, initially cloudy, later half overcast; during
the afternoon distant thunder showers followed by half to full overcast;
29 July — heavily overcast, somewhat less during middle of day, some sunshine
later followed by heavy overcast and drops of rain; moderate N-NE wind;
30 July — heavy overcast before 8:00 a.m., light to heavy overcast throughout
day, light N wind;
1 Aug. — initially cloudy, completely overcast between 8:30 and 10:00 a.m.,
followed by some variation in light NE wind;
5 Aug. — sunny, warm weather, some distant clouds.
The general picture of the diurnal activities can be sketched as follows. In the
morning before 8:30 in sunny weather few or no insects are seen on the open
sandy soil. Initially there is also no activity in the Salix growth. As the effect of
the sun becomes stronger, the latter places begin to show signs of life, soft
stridulating is heard and here and there the insects begin to eat the grasses.
Males soon arrive on the sand; they jump rapidly out of the plant growth,
sometimes two or three at once, and then return just as rapidly. Once in a while
a female is seen, often pursued by a number of males. In the shrub stridulating
and courting increase. After 9:00 a.m. there are more insects on the sand. Most
of the females are animals which, in view of the size of their abdomens, will
shortly begin to oviposit. These females often are seen digging into the sand with
their abdomens. Now and then this digging is followed by oviposition. The laying
of eggs is confirmed by the size of the females’ abdomens. The females then
usually disappear again into the Salix or along its edges (oviposition in the shrub
was not observed).
As soon as the sun is covered by clouds the male song ceases and the number
of arrivals from the Salix on the sand also drops sharply.
In the middle of the day when the weather was very warm, the activity was
concentrated under the Salix overhanging the sand. After 6:00 p.m. most of the
insects return to the shrub, where they remain throughout the night.
On days when the sky was completely overcast (e.g., 29 July, 1955), few insects
were seen on the sand. When the sun came out briefly during the afternoon or a
little radiation penetrated the thin cloud cover, more insects appeared on the sand
although activity remained low. Although oviposition was primarily observed on
sunny days, egg-laying females were also seen on darker days when a number of
days of dark, cold weather were followed by a day with higher temperatures. This
is undoubtedly a case of increased drive, as a result of which oviposition is resumed
even under less favourable conditions. During the six observation days, boring was
seen 161 times, followed in 40 cases by oviposition.
It should be noted that the observations included in Table 44 do not directly
reflect the movements from and to the surrounding vegetation; the course of the
activities is presented as a static picture. Table 45, however, gives the numbers of
adult insects, which in each of the half-hour observation periods on 30 July, 1955,
went from the Salix and type IV vegetation to the sand and back. This table clearly
shows that Table 44 gives an inaccurate picture of the local situation: the actual
population is in motion, and it is quite clear that the presence of the denser
vegetations in the immediate vicinity is of conclusive significance.
B. M. LENSINK: Distributional ecology of Acrididae 433
Table 45. Observations on activity of M. maculatus.
From Salix-shrub | From bare area From Type IV From bare area
to bare area to Salix-shrub to bare area to Type IV
7:30— 8:00 — — — —
8:00— 8:30 1 — — —
8:30— 9:00 3 — = —
9:00— 9:30 10 — 2 —
9:30—10:00 4 3 1 —
10:00—10:30 2 — = 2
10:30—11:00 7 2 4 2
11:00—11:30 3 5 1 —
11:30— 12:00 6 2 2 1
12:00—12:30 6 4 4 4
12:30— 13:00 7 U 2 1
13:00—13:30 7 11 1 1
13:30—14:00 4 1 3 1
14:00—14:30 5 3 3 4
14:30—15:00 12 4 5 2
15:00—15:30 5 4 1 —
15:30— 16:00 2 1 6 3
16:00— 16:30 2 3 5 =
16:30—17:00 5 — 2 —
17:00—17:30 4 6 4 1
17:30—18:00 5) 3 — 1
18:00—18:30 2 4 => crm
A simular situation is found in the part of the terrain in which Quadrats 2
and C are located, where the behaviour of M. maculatus during the day was also
studied. During sampling, this species was seen regularly. Closer observation
showed that they came from the neighbouring vegetation during the morning and
returned there during the evening. The activity was often great: there was a
continuous procession from the bordering vegetation to the open vegetation and
back again.
Concerning the factor of food, it may be noted that from among the large num-
bers of species of herbs and grasses, there is a strong preference for the grasses.
Observations concerning the use of food plants were made during the study of the
diurnal activities in particular, but also during other investigations. A list is given
in Table 46 of a number of plant species predominating in the vegetation of the
terrain studied, and the numbers of times M. maculatus was observed feeding on
these plants is indicated.
The preference for grasses and Carex arenaria is clear, and Festuca rubra subvar.
arenaria forms the main food of the grasshoppers under observation. It was striking
that nymphs and adults preferred blades of grass lying on the ground to erect ones.
Although especially the tips of the blades are gnawed off and eaten, grasshoppers
were seldom seen in the erect grass. The wide, stiff leaves of Calamagrostis
epigejos are also seldom used for climbing. The first instar fed at their hatching
sites (types I and II), especially on Phleum arenarium which is abundant during
434 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
the hatching period (April and May), and on Festuca rubra subvar. arenaria. It is
striking that almost no herbs are consumed. When the insects have no other choice,
for instance in the laboratory, herbs are also used as food. Galium verum var.
maritimum is, however, consistently refused, and when a grasshopper gnaws one
of its leaves, death follows rapidly. In this connection it is remarkable that for
O. coerulescens it was observed several times in the field that these insects consume
Galium verum var. maritimum without adverse effects as shown by the check.
The diurnal activities of C. biguttulus
The low population density of C. biguttulus in the terrain meant that only a
small number of observations of the activities of this species could be collected.
The study of the distribution of C. biguttulus indicated that in the period in which
older nymphs and adults occur, they are to be found primarily ín the dense grass
cover (types V and VI). Most of the insects were found, however, in the grassy
edges of the south side of the hawthorn hedges surrounding the terrain, but for
technical reasons these grassy strips could not be quantitatively sampled. The
observations of the activities of these insects were done mainly in these border
strips, although they were greatly hampered by the thickness of the grasses.
Most of the diurnal activities of C. biguttulus, as for M. maculatus, were ob-
served on warm, and especially sunny, days. The insects are unusually active under
these conditions, particularly the adult males, which walk through the vegetation
a great deal. In addition, and more than for the other two species, they are observed
to make jumps for which they sometimes use their wings. In a few cases, males
could be followed over long distances thanks to an aberration in song. In this way
it was observed that on warm and sunny days they covered a distance of about
20 metres through various types of vegetation within a period of three hours,
without using the wings in jumping. C. brunneus also shows this tendency to cover
long distances often without flying. On a few occasions it was observed that an
adult flew from a sun-warmed wall to a height of 50—100 cm above the ground
and landed 10 to 15 metres away in a hedge, from which a new flight soon
followed.
During the morning many of the insects sit sunning themselves. For this they
often use wide leaves, particularly horizontal ones like those of blackberry bushes
and the lowest branches of the hawthorn trees. With strong solar radiation they
become more active and alternate between sitting in the full sunshine and in the
shade provided by leaves and stalks.
A special form of migration was observed in part of the terrain shaded on its
southern side in the afternoons by a wide, high hawthorn hedge. The vegetation
consisted of a dense grass cover (type VI) shifting in the direction away from the
hedge to a type V vegetation. In the morning, adults of C. biguttulus were seen
in type VI vegetation. In the afternoon, when this vegetation was in the shade,
the largest numbers of insects were to be found in the type V, which still lay in
the sun. Observations at this place showed that as the shade extended over the
vegetation, the insects avoided it as much as possible by shifting to sunny spots.
This occurred for the most part by walking, but also with small jumps. This
B. M. LENSINK : Distributional ecology of Acrididae 435
species is distinguished in its behaviour by being markedly more active than the
other two species, which enables it to find the spots with the most sun.
The diurnal activities of C. albomarginatus
The activity of C. albomarginatus was studied in the most heavily populated
parts of the terrain: the Calamagrostis epigejos growths which included Quadrats
6, F, and G. In this vegetation, observation is difficult because the grass is so thick
and the grasshoppers can be seen only along the edges of the growth.
The activities were nevertheless studied within the vegetation, making use of a
ladder placed over the grass. The series sampling showed clearly that in 1955 in
this growth few insects were present, i.e., one to nine insects per square metre
(on the sampled quadrats, while along the edges the number was estimated at
at least fifteen per square metre).
The broad picture sketched for M. maculatus agrees in general with that for
C. albomarginatus. The activity is highest with favourable weather conditions; in
dark or rainy weather little movement is observed. Continuous observations were
made in sunny weather in July and August, 1955 and give a picture of the insects
activities in this vegetation during the day. Quantitative data are almost impossible
to collect, however, because of the relatively few insects seen.
Before 9:00 a.m. little occurs in the grass cover. Most of the insects are found
at a level just under the upper grass leaves at a height of 40—60 cm, which
particularly in the middle form a dome above the space between the leaves and
stems. Between 10:00 and 11:00 a.m. on the sunny side (SE) many insects were
seen between the blades at a height of 10—20 cm above the ground. They sit with
their heads pointing upwards sunning themselves, a few eating from the leaf.
They are also observed on dead grass on the ground, and in the surrounding, less
dense vegetation.
On the shady side (NW), however, none are found on the ground; they sit in
the tops of the grass, sunning themselves and eating. Later in the morning courting
occurs, but on the whole these insects are not very active. There is much less
movement than is seen for M. maculatus.
The border areas which are shaded in the morning, now begin to receive more
sun, and the insects move towards lower levels while in the now shaded parts
most of them seek higher places between the grass blades. At about 3:00 p.m.
the picture is the opposite of the morning situation. After 4:00 p.m. it is observed
that on the side of the vegetation on which the sun shines, the nymphs and
adults begin to climb higher in the vegetation. When the sun sinks lower, the
leaves and stalks of the surrounding vegetation create a great deal of shade, and
this probably explains the vertical migration. The grasshoppers which find them-
selves shaded by a leaf climb continually higher, consistently seeking the sunniest
spots. When there is a lot of wind on the sunny side, most of the insects sit to sun
themselves not on the outer side, but between the first rows of blades. Later in the
day courting drops in intensity and more eating insects are seen. From this point
activity decreases and the largest numbers are found high between the stalks.
The consistent search for the ’’sun’’ was successfully observed in an open space
436 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
with a diameter of about 75 cm in the middle of the vegetation. The largest
numbers were at first found high between the leaves, and then as the sun rose in
the sky they went lower on the side exposed to the sun, and continued to choose
the sunny side until later in the afternoon they were again found high between
the tops of the plants.
The reactions to various weather conditions indicate that this species, too, is
extremely sensitive to heat, and especially that due to direct radiation. As STOUT-
JESDIJK (1961) demonstrates, in dense grass vegetations like the Calamagrostis
growths mentioned here, the highest temperatures are measured at the level of the
leaf-tips, and this is where during the entire day, in the summer months, the
nymphs and adults of C. albomarginatus were observed. They migrate to lower
levels only when there is a great deal of wind.
In irregular, dense vegetations the highest temperatures are measured in the
layer of dead leaves on the ground. A comparable situation is encountered in our
case, namely in the opening in the dense vegetation and along its edges where the
dense grass cover is very irregular and shows a gradual transition to more open
vegetation. Where the rays of the sun can penetrate to the soil, the highest tem-
peratures are found on the ground. When this radiation is interfered with by
shade from the surrounding dense grass cover, the highest temperatures are found
at higher levels where the direct effect of the radiation is found. The nymphs and
adults of C. albomarginatus consistently seek the warmest places in the grass cover,
producing vertical migration along the edges and in open spaces of the dense
grass cover.
For this species a few data were also collected concerning its choice of food in
the field. The dense, high grass cover in which most of the observations concerning
diurnal activity were made, is composed of a vegetation of Festuca rubra subvar.
arenaria and Carex arenaria, dominated by Calamagrostis epigejos. Other grasses
and herbs are poorly represented, including Holcus lanatus, Poa pratensis var.
humilis, Achillea millefolium, Plantago lanceolatum var. sphaerostachya, Vicia
cracca and Rumex acetosa. The 41 observations of eating habits in this vegetation
are shown in Table 47.
The first two grasses, which form the principal mass, are the most frequently
eaten.
In 1956 a number of nymphs were put in a cage which was placed over a piece
of grass cover of type V vegetation. In this grass cover the predominating species
were Festuca rubra subvar. arenaria and Agrostis tenuis, and Calamagrostis ept-
gejos, and in lesser amounts Carex arenaria, Galium verum var. maritimum, Achil-
lea millefolium, Poa pratensis var. humilis, Cerastium arvense, Thymus pulegioides,
and Lotus corniculatus var. ciliatus. Twenty nymphs of the second to the fourth
instar were observed within two hours; a total of 93 feeding observations were
made (Table 48).
In this type of vegetation, too, only grasses were eaten, herbs were not touched.
A similar arrangement was used over a grass cover of type IV vegetation in
which the herbs predominated more than in the above-mentioned cover. Forty
feeding observations were made in two hours of observing (Table 49).
Here again the grasses are preferred. Calamagrostis epigejos was not eaten in
B. M. LENSINK : Distributional ecology of Acrididae 437
Tables 46—49. Numbers of feeding observations, for explanation see text.
Table 46 M. maculatus Table 47 C. albomarginatus
Festuca rubra subvar. arenaria 261 Festuca rubra subvar. arenaria 23
Festuca ovina 19 Calamagrostis epigejos 16
Calamagrostis epigejos 7 Carex arenaria 1
Phleum arenarium 1 Vicia cracca 1
Poa pratensis var. humilis 0 —
Agrostis tenuis 13 No. of feeding observations 41
Corynephorus canescens 0
Carex arenaria 28 Table 48 C. albomarginatus
Galium verum var. maritimum 0 Festuca rubra subvar. arenaria 38
Lotus corniculatus var. ciliatus 0 Agrostis tenuis 13
Thymus pulegioides 0 Calamagrostis epigejos 37
Leontodon nudicaulis 0 Poa pratensis var. humilis 5
Hieracium pilosella 0 ==
Achillea millefolium 1 No. of feeding observations 93
Cerastium arvense 3
Table 49 C. albomarginatus
No. of feeding observations 333 Festuca rubra subvar. arenaria 17
Agrostis tenuis 13
Festuca ovina 3
Carex arenaria 4
Plantago lanceolata var. sphaerostachya 3
Calamagrostis epigejos 0
No. of feeding observations 40
the observation period, but is very thinly distributed with few stalks in this cover
and was therefore rarely encountered.
When the insects have a free choice, their food is chiefly the grasses and Carex
arenaria. These grasses occur in almost the entire terrain. Herbs are apparently
not considered as food even when they occur frequently in the vegetation. In the
laboratory such herbs as Lotus corniculatus var. ciliatus, Achillea millefolium,
Hieracium pilosella, and Thymus pulegioides are eaten when little or no grass is
present. Gallium verum var. maritimum was consistently refused by C. albomar-
ginatus when offered as food in the laboratory. When an occasional insect
nibbled” on a leaf of this plant, it died within 24 hours.
SUMMARY
(1) Aspects of the lives of three species of grasshoppers (M. maculatus, C.
biguttulus, and C. albomarginatus) in their environment were investigated. These
investigations were intended to provide a better understanding of the distribution
of these species in the dunes of Voorne (The Netherlands). This distribution was
seen as an aspect of the inter-relation of the animated and the inanimate world.
This inter-relation can be considered in terms of the related quantities or in terms
of the phenomenon of the relationship itself, in the sense of interaction between
the factors.
The investigation described here was based on a number of selected quantities:
the development of the vegetation with the change of the microclimate and the
438 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
development of the grasshoppers. A study was made of the influences which the
inter-relationship of the quantities exercises on the distribution of the grasshoppers.
This inter-relationship has a special expression in the occurrence of migration in
many phases of the life of the insect, another indication that this inter-relationship
is dynamic in nature.
(2) The technique consisted of a series of samplings in various types of
vegetation distinguished according to structure. Each sampling reflects the situation
at a given time. A series of these samplings throughout the year gives a picture of
the modifications occurring in the distribution of the grasshopper population. In
addition to the investigation of the distribution of the various immature stages of
the insects, attention was also given to their activities during the day under various
weather conditions and the influence of the microclimate. The essential points may
be recapitulated here.
(3) The vegetation is for the grasshopper one of the most important and
predominant factors in its environment. Variations in distribution, however, cannot
be explained on the basis of dependence on the vegetation as food source because
in the dune region the insects eat many species of plants and certainly the most
frequently occurring species of the grasses. What the investigation showed especi-
ally clearly was that the structure of the vegetation plays an important part in the
distribution observed. It was also seen that the characterization of the structure
of the vegetation by means of the French-Swiss method of vegetation description
was too limited. Seven structural types had to be distinguished in the present study.
These types and the internal changes occurring in the course of the year and from
year to year are described.
The various types of vegetation have a strong influence on the microclimate.
Since both the vegetational structure and the macroclimate are subject to changes
in the course of time, the microclimate will also vary continually.
(4) The distribution of the grasshoppers is determined in the first instance
by the site of oviposition. The location of these sites was studied by field sampling
during the time in which the nymphs hatch. This study showed that various influ-
ences are present, such as the distribution of the adults, and the choice of ovi-
position substrate in the preceding year. These influences and the influence of
the combination of vegetation and microclimate on the egg, were expressed not
only in the peculiar distribution of the hatching sites but also in the time at which
the nymphs appeared in the various types of vegetation and the numbers of nymphs
hatched there.
(5) The further distribution of the insects from the hatching sites over the
field is distinctly affected by the development of the vegetation and events in the
microclimate, which close interconnection is striking.
In general, it may be said that from spring to summer migration occurs from
the less dense to the denser vegetations, a tendency observable in all three species.
Concerning two aspects, the microclimate and the demands made by the insects
in the various immature stages on their surroundings, the following may be said.
The first instars of M. maculatus and C. biguttulus are almost unable to find
shelter against strong solar radiation, hail storms, etc., in vegetation types I and II.
They migrate and we then see an increase in the numbers of the third and fourth
B. M. LENSINK: Distributional ecology of Acrididae 439
instar in type V. In type V the microclimate is more clement and high temperatures
do not occur; but the fertilized females require an oviposition substrate, causing
a return to types I and II. In types III and IV, in which structural changes are less
distinct, the insects find not only shelter in the shade of grass tussocks and between
the herbs covering the ground, but also adequate opportunity to sun themselves.
They are able to maintain themselves in this type of vegetation.
Nymphs of C. albomarginatus, hatched in types III and IV, leave this vegetation.
At this time an increase in this species is seen in type VI.
(6) As an example of the influence of the macroclimate, the following may
serve: As a result of the cloudy weather in the spring of 1955, the microclima-
tological conditions on plots with vegetation of types I and II were far less extreme
than in the preceding year. We see in consequence that the insects remained
much longer on these plots, another indication that these conditions are the primary
factor under the influence of which the nymphs leave this type of terrain, and not,
for instance, the food factor. The climatological conditions had an inhibitory
effect on migration, but an activating effect on the growth of the vegetation.
Consequently all types of vegetation were denser at the time of migration than in
both preceding years, so that migration was more restricted than in 1953 and 1954.
During both these years, only the edges of type V were populated by M. maculatus
and C. biguttulus. This vegetation approached more closely the characteristics of
type VI. C. albomarginatus was also found only in the border zone of type VI.
It may be concluded from this that the insects inhabit the same types of vege-
tation as in the preceding year, but their distribution over the inhabited terrain
varies, and may do so from year to year.
(7) During the summer months the situation is as follows: M. maculatus and
C. biguttulus are found seldom or never in types I and II, in small numbers in
types III and IV, and in the largest quantities in type V. C. albomarginatus is
found especially in type VI or in the dense grass borders of type VII; elsewhere
the numbers are small. In this period the vegetation has reached its maximum
growth, and it is thanks to the many horizontal leaves in type V that a ’’geophilic’’
insect like M. maculatus (and this holds to a certain extent for C. biguttulus as
well) can live in this dense grass cover. In the dense, high grass cover, indeed,
we find these species almost exclusively along the edges, while C. albomarginatus
is able to move throughout this heavy cover.
Local variations from this situation are found in the distribution of M. maculatus
and C. biguttulus. In a few spots covered with types I and II vegetation, individual
specimens of these species were seen throughout the season, first nymphs of the
various stages and later adults as well. This was possible because of the heavy
grasshopper traffic between these terrains and the surrounding types V and VII
vegetations. Observations during the day showed that on warm days many of the
females search for a suitable oviposition site, which is rarely or never found in
type V, and during this time they are pursued by males. These activities, however,
are strongly dependent on the weather conditions. In dark, cold weather or in the
early hours of the morning or evening, most of the insects are to be found in the
denser grass cover. Only with increasing radiation does the activity increase and
a varying number moves in and out of the vegetation. But also within a given
440 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
type migration can be observed throughout the day. With strong sunshine it is
seen that the insects make alternate use of sunny and of shaded, cooler spots.
As a result, we see that in their area the greatest numbers of individuals of
M. maculatus and C. bigattulus are found where not only suitable oviposition sites
but also suitable places in which to live occur in the immediate vicinity. In parts
of the terrain comprising many square metres of suitable oviposition substrate,
the population density is very low if no suitable places for the nymphs and adults
are available in the neighbourhood. Where suitable type V places in which to live
are found, M. maculatus is encountered only in places which border on an ovi-
position terrain. C. biguttulus is found much deeper in these growths; it is also
far more active than M. maculatus.
(8) Later in the season, from the middle of August on, new changes take place
in the distribution. In the open vegetations, increasing numbers of adults are
observed, and in the denser types V and VI there is a distinct decrease (for C.
albomarginatus) or even total disappearance (the other two species). In the first
place, it seems self-evident to assume that with increasing age the adults make
different demands on the environment (oviposition). In the second place, changes
in the microclimate in the late summer period begin to play an important part.
The continually lower position of the sun and the cooler nights accompanied by
local mist clearly affect the denser vegetation in which the day and night tem-
peratures are both lower than in July. In the more open vegetations the soil still
reaches appreciable temperatures during the day and even during the night remains
warmer than in the denser vegetation. In this period females are regularly seen
almost ready for oviposition. They are then found in places where they have not
been seen previously, as for instance in types I and II. The distribution of these
females is a contributory factor for the distribution of the young insects in the
spring. The late summer migration, however, is not so extensive that adults are
found in all parts of the terrain with open vegetation. During October the last
of the adults die.
DISCUSSION
(1) The preceding summary was given in order to bring out the peculiar
features contributing to the picture of the distribution of the grasshoppers in a
given area. This basis requires a still broader interpretation to include all these
phenomena as a whole. The problem is to determine in what framework to place
the picture which evolved from the investigation.
In discussing the relationship between organism and environment, I would like
to take as my starting point the concept habitat’ as one of the frequently used
ecological concepts. It is used by many investigators, but always formulated some-
what differently. The difficulty arises from the fact that there is a constant shift
in emphasis from animal to environment and from environment to animal, while
the interrelation between the two is often left very vague. Some authors use a
‘’physiognomic areal concept” (see CARPENTER, 1956) in which the habitat is
defined in terms of the character of the landscape or on the basis of an ‘’environ-
mental factors concept” which includes only selected factors (see CARPENTER,
B. M. LENSINK : Distributional ecology of Acrididae 441
1956) or, as proposed by CLEMENTS (1905): the habitat is “the exact equivalent
of the term environment”. PEARSE (1926) includes animal and environment in his
definition of habitat: there is some place where an animal system of activities
finds an environmental system of activities’. PARK (ALLEE et al., 1949) makes
the relationship of the animal with its environment clearer by pointing out that
the species of animal in its habitat has reached a "survival level”. In its environ-
ment it has become ’’self-sustaining’”. The various definitions contain elements
indicating that there is a tendency to place the inter-relationship between animal
and environment more in the foreground. PEARSE, for instance, mentions the pos-
sibility of the animal to maintain "a reasonable condition of stability” in its
existence. In this phrase, however, the inter-relationship is more assumed than
made the keystone of the definition.
Prus (1954) and BODENHEIMER (1958) see the relationship more sharply.
PEUS proposes the concept ‘’Oekologischer Umwelt’ and BODENHEIMER more or
less in agreement with PEUS, uses the concept ”Ecoworld” to emphasize the con-
nection somewhat more strongly. BODENHEIMER understands under this term: ’’the
combination of all exogenous and endogenous factors, processes and organisms
which have any relation — whether directly or not directly perceived — to the
living species’. And to avoid any misunderstanding of his meaning, he again
clearly states: The species itself is of course an integral part of this eco-world”.
Although not expressed directly, it is implied here that BODENHEIMER makes the
integration of the quantities so central that the topographical aspect is neglected in
his approach.
(2) The observations indicate that a study such as this cannot be limited to
the simple notation of the places in which the animals occur, a technique which
results in what is called a pattern or mosaic of the distribution. The actual situation
in the field requires a somewhat different basis. The presence of the animal cannot
be seen as simple presence, it must be seen as the representation of a situation
belonging to a particular moment in a particular place. This approach introduces
the character of a transitional situation, ie, it assumes its significance when
considered in the light of what has already occurred and what is to occur.
In addition to changes in distribution from year to year and within each year,
from week to week and from day to day, there is also variation within each day
which is closely related to the development of the vegetation and the microclimate
and the physiological condition of the animal. The situation at the end of one
year also determines the situation at the beginning of the next.
The complexity of the inter-relationship becomes clear as soon as the external
factors seem to predominate in determining the course of events, as for instance in
spring when the quantity of solar radiation is important for the hatching of the
eggs, or when the extreme conditions of terrains of type I and type II are primarily
responsible for migration to the denser vegetations. Then again it is the internal
factors that are of dominant importance, for instance because the fertilized females
have specific substrate requirements to lay their eggs or because the distribution
within the vegetation is limited because of the nature of the behaviour or structure
of the organs or locomotion.
It is clear from the foregoing that the distribution itself has a developmental
442 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
process, for which a complex inter-relationship between internal and external fac-
tors is responsible. In the distribution of the insect, the nature and result of this
interaction is expressed, which is to say that it provides the basis for drawing
conclusions about the inter-relationship.
The concept habitat’ discussed above is inadequate to cover the inter-relation-
ship, which is both functional and topographical in nature. To express both aspects
in the inter-relationship more effectively, I would like to propose the concept ’’vital
space”. The “vital space” is the totality of relationships in which animal and
environment are involved with each other, and one which is subject to change from
moment to moment. We have to do with a complex of relationships between
quantities in which the relationships are expressed in “actions”. These actions”
can be deduced from aspects of the activities of the animals, and this describes the
character of their vital space.
ACKNOWLEDGEMENTS
I wish to express my sincere gratitude to Prof. Dr. D. J. Kuenen for his
encouragement and stimulating criticism during the completion of this work. I also
wish to extend my thanks to Mr. J. J. Busch for the invaluable stimulation of his
sustained interest in my work, and to Mr. E. van der Maarel, Dr. P. Smit, and
Dr. Ph. Stoutjesdijk for their productive criticism. I am grateful to Mrs. I. Seeger
for the translation of this paper in which its original character is retained, to
Mr. J. G. M. Marquenie for the photographs, and Mr. N. Teegelaar for the
preparation of the figures.
I am greatly indebted to The Netherlands Organization for Pure Scientific
Research (Z.W.O.) for the suppott which made this field study possible. I wish
to express my appreciation to the Board of the Foundation for Scientific Duneland
Research for the hospitality extended to my wife and myself at the Weevers’ Duin
Biological Station at Oostvoorne between 1953 and 1955. I am also obliged to
the Uyttenboogaart-Eliasen Stichting for a grant that made it possible to appoint
an assistant in this work for six months.
The Nederlandsche Entomologische Vereeniging kindly accepted this paper for
publication.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963 PLAAT
Pl. 2. Landward dunes, the area of investigation
B. M. LENSINK : Distributional ecology of Acrididae
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963 PLAAT 3
PI. 3. Vegetation types I, II, and V; the light area is type I, surrounded by type II, and at
right rear is the border of type V. The hawthorn hedge can be seen in the background
B. M. LENSINK : Distributional ecology of Acrididae
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963 PLAAT 4
Pl. 4. Vegetation type III
B. M. LENSI
K: Distributional ecology of Acrididae
PLAAT 5
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 106, AFL. 8, 1963
ion type VI
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B M
ENTOMOLOGISCHE BERICHTEN
De Entomologische Berichten worden eveneens door de Nederlandsche Entomo-
logische Vereeniging uitgegeven en zijn bestemd voor de publicatie van kortere
artikelen, van faunistische notities etc, alsmede van de Verslagen der Vergade-
ringen en van mededelingen van het Bestuur. Zij verschijnen twaalf maal per jaar
in een aflevering van 16 of meer bladzijden. Deze 12 afleveringen vormen samen
een deel.
TRICHOPTERORUM CATALOGUS
Ook de Trichopterorum Catalogus van F. C. J. Fischer wordt door de Neder-
landsche Entomologische Vereeniging voor rekening van de auteur uitgegeven.
De eerste vier delen verschenen in 1960, 1961, 1962 en 1963. Er zullen nog elf
delen volgen, in totaal worden het er 15. Alle bekende recente en fossiele soorten
van de gehele wereld met hun synoniemen worden er in behandeld. Van alle
genera worden de type-soorten aangewezen en de literatuur zal van 1758 tot eind
1960 met de geografische verspreiding vermeld worden. Delen I en II kosten
f 39,— per stuk, delen III en IV f 45, —. Van alle delen zijn ook éénzijdig bedrukte
exemplaren beschikbaar. De prijsverhoging hiervoor bedraagt f 2,50.
Alle zakelijke correspondentie betreffende de Vereeniging te richten aan de
Secretaris, W. Hellinga, Weesperzijde 2311, Amsterdam-O.
Alle correspondentie over de redactie van het Tijdschrift voor Entomologie te
richten aan de Horn Dr. A. Diakonoff, Rijksmuseum van Natuurlijke
Historie, Leiden.
Alle correspondentie over de redactie van de Entomologische Berichten te rich-
ten aan de Redacteur, B. J. Lempke, Oude IJselstraat 12111, Amsterdam-Z. 2.
Alle betalingen te richten aan de Penningmeester, Drs. H. Wiering, Doorntjes
29, Bergen (N.H.), postgiro 188130, ten name van de Nederlandsche Entomolo-
gische Vereeniging te Bergen (N.H.).
Alle correspondentie betreffende de Bibliotheek der Vereeniging te richten aan
de Bibliotheek, Zeeburgerdijk 21, Amsterdam-O.
NOTICE TO SUBSCRIBERS
All correspondence referring to subscriptions and exchange of the periodicals
of the Netherlands Entomological Society should be addressed to the Librarian,
Zeeburgerdijk 21, Amsterdam-O, Netherlands.
NOTICE TO CONTRIBUTORS
Contributors will receive free of charge fifty reprints of their papers, joint authors have
to divide this number between them at their discretion. Additional reprints may be ordered
when returning proofs; they will be charged at about two Dutch cents per page.
Manuscripts should be written in Dutch, English, French, German or Italian. If they
contain descriptions of new genera, species, etc., they should be in one of the four last
mentioned languages: only when the descriptions form a minor part of the paper, the
manuscript may be written in Dutch, with the descriptions in one of these languages. Papers
in Dutch should contain a short summary in one of these four languages.
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Names of genera and lower systematic categories, new terms and the like are to be under-
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Authors will be charged with costs of extra corrections caused by their changing of the
text in the proofs.
Bibliography should not be given in footnotes but compiled in a list at the end of the
paper, styled as follows:
Mosely, M. E., 1932. ”A revision of the European species of the genus Leuctra (Pleco-
ptera)”. Ann. Mag. Nat. Hist., ser. 10, vol. 10, p. 1—41, pl. 1—5, figs. 1—57.
Text references to this list might be made thus:
Mosely (1932) says....’ or ’’ (Mosely, 1932)”.
The editors reserve the right to adjust style to certain standards of uniformity.
Manuscripts and all communications concerning editorial matters should be sent to
Dr. A. DIAKONOFF, Rijksmuseum van Natuurlijke Historic, Leiden, Netherlands.
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ERNST MAYR LIBRARY