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TIJDSCHRIFT "FR
VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING
| Tijdschrift voor Entomologie, deel 132, 1989
NEDERLANDSE ENTOMOLOGISCHE VERENIGING
BESTUUR (BOARD)
Vooszitter(Chalrma n) Pee J. Krikken
Vice-voorzitter (Vice-President) ................. S. A. Ulenberg
SECIETANISI(SE CE TAG) EE M. J. Sommeijer
AANEEN TRAI en ler Aus Afdeling Sociale insekten RUU,
Postbus 80086, 3508 TB Utrecht
le Penningmeester (Treasurer I) ................. L. P. S. van der Geest
AGUSTA Doornenburg 9, 1211 GP Landsmeer
2e Penningmeester (Treasurer Il) ................ A. P.J. A. Teunissen
AGG RAI NN ENE eS ERE Strausslaan 6, 5251 HG Vlijmen
Bibliochecarisk(llibrarian ee J. P. Duffels
AA DATE RIO A een RIS III ORE FIANO Plantage Middenlaan 64, 1018 DH Amsterdam
IHds(Memben)E EN C. J. Zwakhals
TIJDSCHRIFT VOOR ENTOMOLOGIE
Redaetiel(EditorialiBoard) ER Re e ESE E. J. van Nieukerken, J. van Tol, C. van Achterberg
and S. A. Ulenberg
DALAT EN RUI E AO A oa Ryksmuseum van Natuurlijke Historie, Postbus
9517, 2300 RA Leiden
Samenstelling van de index
(COMPILATOMOMNAEX) Pre R. T. Simon Thomas
The journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea.
Subscription rate: DFI. 300.— per year.
Volume 132 appeared in two issues on 3.vii and 1.xii.1989
%
ISSN 0040-7496
INHOUD (CONTENTS)
Askew, R. R., G. G. Cleland, D. A. L. Davies & T. W. Harman. — A report on a collection
of @donata from North Sulawesi Indonesia... e e NS)
Assem, J. van den & M. J. Gijswijt. — The taxonomic position of the Pachyneurini
(Chalcidoidea, Pteromalidae) as judged by characteristics of courtship
BEVONO III 149
Aukema, B. — Annotated checklist of Hemiptera — Heteroptera of The Netherlands 1
Baldizzone, G. — A taxonomic review of the Coleophoridae (Lepidoptera) of China.
Contribution to the knowledge of the Coleophoridae, LH ............. 199
Batten, R. — Two new Mordellidae (Coleoptera) from Kashmir and Sumatra ........ 105
Belle, J. — Phyllogomphoides indicatrix, a new dragonfly from Mexico (Odonata: Gom-
Pi AN e NEO ere one I 155
Belle, J. — Epigomphus corniculatus, a new dragonfly from Costa Rica (Odonata: Gom-
PRIORA LARE ae an et eee 158
Belle, J. — A revision of the New World genus Newraeschna Hagen, 1867 (Odonata:
ASIA) ET 259)
Blackith, R. E., see Butlin, R. K.
Blackith, R. M., see Butlin, R. K.
Brooks, S. J. — New dragonflies (Odonata) from Costa Rica .................. 163
Butlin, R. K., R. E. Blackith & R. M. Blackith. — Eumastacidae (Orthoptera) from the
Project Wallace Expedition to Sulawesi (Indonesia) ................ 109
Cleland, G. G., see Askew, R. R.
Davies, D. A. L., see Askew, R. R.
Duffels, J. P — The Sulawesi genus Brachylobopyga (Homoptera: Cicadidae) ........ 123
Duijm, M., see Landman, W., and see Oudman, L.
Gijswijt, M. J., see Assem, J. van den.
Harman, T. W., see Askew, R. R.
Heijerman, Th. & H. Turin. — Carabid fauna of some types of forest in The Netherlands
(@oleoptera:|Catabidac)y ee LIES o MM I ee 241
Helsdingen, P. J. van & E. J. van Nieukerken. — Alexey Diakonoff, 1907-1989 ....... 161
Kanaar, P. — A new Saprinus from Irian Jaya (Coleoptera: Histeridae)........... 285
Landman, W., L. Oudman & M. Duijm. — Allozymic and morphological variation in
Ephippiger terrestris (Yersin, 1854) (Insecta, Orthoptera, Tettigonoidea) 183
Landman, W., see Oudman, L.
Oudman, L., W. Landman & M. Duijm. — Genetic distance in the genus Ephippiger
(Orthoptera, Tettigonoidea). — a reconnaissance . . ................ 1157074
Oudman, L., see Landman, W.
Platel, Th. G. H. — The egg laying and larval development of Pholcus phalangioides (Fuesslin)
(AganedesPholeidae iia, sue eee TE RE IN 135, 288
Rossem, G. van. — The genus Cryptus Fabricius, 1804 of the Canary Islands (Hymenoptera,
Tél UN O HITS) tes aan RO I O RI E CI CIRCO Dl
Turin, H., see Heijerman, Th.
Wittmer, W. — Die Gattung Polemiosilis Pic auf Sulawesi (Coleoptera, Cantharidae). (41.
Beitrag zur Kenntnis der indo-malaiischen Cantharidae) ............. 129
Booksreyiewsiand announcements 1". NN NU 114, 122, 128, 162, 182
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CRI AGI RET 44 Pi
TRO
DER? AFLEVERING 1 1989
TIJDSCHRIFT
VOOR ENTOMOLOGIE
UITGEGEVEN DOOR
DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING
NC
LIBRARY
n 99 1020
AUG Li 2 1999
HARVA
UNIVER EEY
Tijdschrift voor Entomologie
A journal of systematic and evolutionary entomology since 1858
Editors: E. J. van Nieukerken and J. van Tol
Co-editors: C. van Achterberg and Mrs. S. A. Ulenberg.
Advisory board: M. Brancucci (Basel), Y. Hirashima (Fukuoka) and M. R. Wilson (London).
The “Tijdschrift voor Entomologie” is published in two issues per year by the "Nederlandse Entomologische
Vereniging” (Netherlands Entomological Society), Amsterdam.
Editorial address: c/o Rijksmuseum van Natuurlijke Historie, Postbus 9517, 2300 RA Leiden, The Nether-
lands.
All enquiries of membership of the Society, as well as correspondence regarding subscription and possibilities
for exchange of this journal, should be addressed to
Nederlandse Entomologische Vereniging,
c/o Instituut voor Taxonomische Zoologie,
Plantage Middenlaan 64,
1018 DH Amsterdam
The Netherlands
Subscription price per volume for non-members: Hfl. 300.—
INHOUD CONTENTS
B. Aukema, Annotated checklist of Hemiptera-Heteroptera of the Netherlands.........
R. Batten, Two new Mordellidae (Coleoptera) from Kashmir and Sumatra............
R. K. Butlin, R. E. Blackith & R. M. Blackith, Eumastacidae (Orthoptera) from the Project
Wallace ExpeditronitoSulawesi{(Indonesia) 2... e ER E eee
BOokanNOUNEEMENESTANAEVIEWSERRR ER O EE
R. R. Askew, C. G. Cleland, D. A. L. Davies & T. W. Harman, A report on a collection of
OdonatatromiNorthisulawestindonesias RER
EFaunaMalesiana Foundation 3. . „er. EAN RI NE
J. P. Duffels, The Sulawesi genus Brachylobopyga (Homoptera: Cicadidae) .............
Book'announcements and reviews..." IE
W. Wittmer, Die Gattung Polemiosilis Pic auf Sulawesi (Coleoptera, Cantharidae).
(41. Beitrag zur Kenntnis der indo-malaiischen Cantharidae) ....................
Th. G. H. Platel, The egg laying and larval development of Pholcus phalangioides (Fuesslin)
(Araneae: Pholcidae) een e LL ce E
J. van den Assem & M. J. Gijswijt, The taxonomic position of the Pachyneurini (Chalci-
doidea, Pteromalidae) as judged by characteristics of courtship behaviour ...........
J. Belle, Phyllogomphoides indicatrix, a new dragonfly from Mexico (Odonata: Gomphi-
ALENIA I NO ate ne PNE LIT RAI EN NEPI
(GTS) ea ent Re e MO A UR Li co ose
Publilshed 3 July 1989
ISSN 0040-7496
ANNOTATED CHECKLIST OF HEMIPTERA-HETEROPTERA
OF THE NETHERLANDS”)
by
BEREND AUKEMA
ABSTRACT
Aukema, B., 1989. Annotated checklist of Hemiptera-Heteroptera of The Netherlands. —
Tijdschrift voor Entomologie 132: 1-104, figs. 1-120, tab. 1. [ISSN 0040-7496]. Published 3 July
1989.
A list of 586 species of Heteroptera recorded from The Netherlands is presented. Local
distribution is indicated by data on the presence in the twelve Dutch provinces, while for new,
rare and endangered species the records are included in more detail. Eighteen species are recorded
for the first time from The Netherlands: Deraeocoris flavilinea (Costa), Tytthus geminus (Flor),
Brachyarthrum limitatum Fieber, Criocoris sulcicornis (Kirschbaum), Reuteria marqueti Puton,
Heterocordylus genistae (Scopoli), Lygus adspersus (Schilling), Lygus wagneri Remane, Lygo-
coris populi Leston, Capsus pilifer Remane, Capsodes sulcatus (Fieber) and Trigonotylus caeles-
tialium (Kirkaldy) (Miridae), Temnostethus longirostris (Horvath) and Scoloposcelis pulchella
(Zetterstedt) (Anthocoridae), Coranus woodroffei Putshkov (Reduviidae), Kleidocerys privignis
(Horváth), K. truncatulus ericae (Horváth) and Scolopostethus pseudograndis Wagner (Lygaei-
dae). Kleidocerys privignis is new for Western Europe.
Key words. - Heteroptera; checklist; faunistics; distribution maps.
Berend Aukema, Pomona 66, Wageningen, The Netherlands.
INTRODUCTION
Between 1853 and 1951 five editions of a check-
list of Dutch Heteroptera appeared (De Graaf &
Snellen van Vollenhoven 1853, De Graaf, Six &
Snellen van Vollenhoven 1860, Snellen van Vollen-
hoven 1868-1878a, 1878, Fokker 1883-1899 and
Reclaire 1932-1951). Because the last supplement
to the list of Reclaire was published in 1951 there
was a growing need for a new updated list.
Reclaire (1932-1951) listed a total of 514 species
from The Netherlands. In the present, sixth list,
586 species are included. Fourteen species of the
list of Reclaire are omitted, because they are either
considered incidentally introduced (Stephanitis py-
rioides (Scott), Pinthaeus sanguinipes (Fabricius)
and Rhaphigaster nebulosa (Poda)), misidentified
(Sigara venusta (Douglas & Scott), Velia rivulorum
(Fabricius), Chartoscirta geminata (Costa),
Orthonotus cylindricollis (Costa), Phytocoris ju-
niperi Frey-Gessner, Xylocoris parvulus (Reuter),
Piesma capitatum (Wolff) and Rhopalus rufus
') Dedicated to the memory of R. H. Cobben, the well-
known Dutch Hemipterist, who died at 7 December
1987.
2) Mededeling EIS-Nederland, nr. 41.
Schilling), or sunk as synonyms (Monosynamma
nigritulum (Zetterstedt), Orthocephalus ferrari
Reuter and Neides favosus Fieber). On the other
hand Microvelia pygmaea (Dufour), Saldula palus-
tris (Douglas), Amblytylus brevicollis Fieber and
Eurydema ventralis Kult had to be added, because
their origin and/or identity, questionable at the
time, could be established. Furthermore, two taxa
previously included as varieties have been raised to
specific status, viz. Drymus rye Douglas & Scott
and Megalonotus sabulicola (Thomson).
The specimens of Coranus tuberculifer Reuter
recorded by Blöte (1954, as f. nov. spec.) belong to
C. woodroffei Putshkov.
Callicorixa wollastoni (Douglas & Scott), Eury-
colpus flaveolus Stal, Phytocoris hirsutulus Flor,
Peritrechus distingendus (Flor), Stictopleurus cras-
sicornis (Linnaeus), Sciocoris umbrinus (Wolff),
Sehirus dubius (Scopoli) and Geotomus punctula-
tus (Costa) are listed erroneously for The Nether-
lands by Stichel (1955-1962).
The recording of Callicorixa producta (Reuter)
from The Netherlands (Polderman & Van der
Velde 1971) was due to misidentification of imma-
ture specimens of Callicorixa praeusta (Fieber);
Jansson (1986) included Micronecta griseola Hor-
N
vath erroneously as a Dutch species by misinter-
preting a locality label (Nieser, in litt.).
Cremnocephalus alpestris Wagner was caught in
Flevoland (7.viii.1982, 1 male, B. van Aartsen, coll.
Museum Amsterdam), but for the present it has to
be considered a casual introduction since it was
caught far away from its natural habitat (pine fo-
rest).
Thus the number of real additions to the list
since 1951 is no less than 80 species (16%). The
following 18 species are recorded here for the first
time from The Netherlands:
Miridae: Deraeocoris flavilinea (Costa), Tytthus
geminus (Flor), Brachyarthrum limitatum Fieber,
Criocoris sulcicornis (Kirschbaum), Reuteria mar-
queti Puton, Heterocordylus genistae (Scopoli), Ly-
gus adspersus (Schilling), Lygus wagneri Remane,
Lygocoris populi Leston, Capsus pilifer Remane,
Capsodes sulcatus (Fieber), Trigonotylus caelesti-
alium (Kirkaldy); Anthocoridae: Temnostethus
longirostris (Horváth), Scoloposcelis pulchella
(Zetterstedt); Reduviidae: Coranus woodroffei
Putshkov; Lygaeidae: Kleidocerys privignus (Hor-
vath), Kleidocerys truncatulus ericae (Horvath)
and Scolopostethus pseudograndis Wagner. [Two
additional species are recorded in an addendum |.
Kleidocerys privignis, thusfar only known from
Bulgaria and Southern Russia (Stichel 1957-1962;
Josifov 1986) is new for Western Europe.
The present list:
2 Pachycoleus waltli
8 Cymatia bonsdorffu
10 Glaenocorisa propinqua
16 Corixa punctata
17-20 Hesperocorixa
18 Hesperocorixa linnaei
21 Paracorixa concinna
22-35 Sigara
23 Sigara stagnalis
24 Sigara hellensti
25 Sigara nigrolineata
35 Sigara lateralis
36 Ilyocoris cimicoides
38 Aphelocheirus aestivalis
43 Notonecta obliqua
51 Velia caprai
55 Microvelia reticulata
59 Gerris lateralis
62-63 Aquarius
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Nomenclature and sequence of taxa above sub-
family level are according to Stys & Kerzhner
(1975) and Stys & Jansson (1988). Below family
level in general the sequence of Southwood & Le-
ston (1964) is followed, with the exception of Co-
rixidae (Jansson 1986), Tingidae (Péricart 1983),
Nabidae (Péricart 1987), Anthocoridae and Cimi-
cidae (Péricart 1972), Lygaeidae (Slater 1964,
1964a), Berytidae (Péricart 1984) and Rhopalidae
(Göllner-Scheiding 1983), which are arranged ac-
cording to the given references. Nomenclature of
Miridae and Lygaeidae in general is according to
the world catalogues of Carvalho (1957-1960) and
Slater (1964) respectively, with regard to grammar
corrections of Steyskal (1973, 1973a). Within
(sub)genera the species are arranged in alphabeti-
cal order. All species are sequentially numbered
throughout. Species with an asterisk (*) before
their number are treated with the same number
under Remarks.
Since the biological and phenological informa-
tion given by Reclaire (1932) is still very valuable,
the changes in names used are listed below to keep
his work accessible. The species numbers used by
Reclaire are those of the catalogue of Oshanin
(1912).
Reclaire (1932):
3236 Pachycoleus rufescens
3475 Cymatia bonsdorffi
3472 Glaenocorisa cavifrons
3405 Corixa geoffroyi
3424-3425 Arctocorisa
3441-3442 Arctocorisa
3425 Arctocorisa linnei
3468 Callicorixa concinna
3413-3423 Arctocorisa
3430-3438 Arctocorisa
3443-3446 Arctocorisa
3413 Arctocorisa lugubris
3423 Arctocorisa hellensi
3446 Arctocorisa fabricu
3419 Arctocorisa hieroglyphica
3364 Naucoris cimicoides
3354 Aphelochirus aestivalis
3400 Notonecta furcata
3281 Velia rivulorum
3283 Velia currens
3279 Microvelia schneideri
3256 Gerris asper
3245-47 Gerris
AUKEMA:
64 Limnoporus rufoscutellatus
69 Halosalda lateralis
70-80 Saldula
85 Campylosteira verna
92 Acalypta platycheila
95 Kalama tricornis
107-109 Physatocheila
109 Physatocheila smreczynskü
111-113 Dictyla
111 Dictyla convergens
113 Dictyla bumuli
114 Agramma laetum
115-117 Loricula
120 Myrmedobia exilis
124-125 Deraeocoris
135 Lopus decolor
138 Conostethus griseus
147 Macrotylus paykulii
149 Orthonotus rufifrons
151-152 Tytthus
157 Plesiodema pinetella
167 Psallus confusus
170 Psallus haematodes
173 Psallus salicis
176 Compsidolon salicellus
177 Phoenicocoris obscurellus
182 Plagiognathus litoralis
192-194 Monosynamma
195 Campylomma annulicorne
192 Monosynamma bohemanni
199 Salicarus roseri
200 Sthenarus rotermundi
202 Hallodapus rufescens
210 Dicyphus pallicornis
225 Orthocephalus coriaceus
229 Fieberocapsus flaveolus
230 Cyllecoris histrionicus
231 Dryophilocoris flavoquadrimaculatus
233 Globiceps fulvicollis cruciatus
239 Heterotoma planicornis
241 Blepharidopterus diaphanus
257 Pseudoloxops coccineus
258 Cyrtorhinus caricis
259 Neomecomma bilineatum
270-272 Orthops
272 Orthops kalmii
273-276 Pinalitus
279-286 Lygocoris
287 Agnocoris reclairei
289 Camptozygum aequale
290-294 Polymerus
295 Charagochilus gyllenhalii
298 Maris striatus
299 Hadrodemus m-flavum
302 Calocoris striatellus
Checklist Hemiptera-Heteroptera
3244 Gerris rufoscutellatus
3298 Haldosalda lateralis
3309-3330 Acanthia
1445 Campylostira verna
1466 Acalypta platychila
1489 Dictyonota tricornis
1602-1604 Physatochila
1604 Physatocheila quadrimaculata
1615-22 Monanthia
1620 Monanthia humuli
1618 Monanthia symphyti
1642 Serenthia laeta
2140-2144 Microphysa
2148 Myrmedobia tenella
2444-2446 Camptobrochis
2824 Onychumenus decolor
2857 Conostethus salinus
2941 Macrotylus paykulli
2956 Byrsoptera rufifrons
2629-2630 Cyrtorrhinus
2972 Plesiodema pinetellum
3017 Psallus diminutus
3030 Psallus roseus
3012 Psallus alnicola
3034 Psallus salicellus
2987 Psallus obscurellus
3095 Plagiognathus fusciloris
3130 Microsynamma
3137 Campylomma annulicornis
3130 Microsynamma bohemani
3157 Sthenarus roseri
3160 Sthenarus rottermundi
2563 Allodapus rufescens
2548 Dicyphus pallidicornis
2755 Orthocephalus mutabilis
2631 Cyrtorrbinus flaveolus
2605 Cyllocoris histrionicus
2607 Cyllocoris flavoquadrimaculatus
2621 Globiceps cruciatus
2687 Heterotoma meriopterum
2654 Orthotylus diaphanus
2686 Pseudoloxops coccinea
2628 Cyrtorrbinus caricis
2639 Orthotylus bilineatus
2403-2405 Lygus
2405 Lygus kalmi
2387-2396 Lygus
2364-75 Lygus
2409 Lygus rubicundus
2414 Camptozygum pinastri
2424-2428 Poeciloscytus
2436 Charagochilus gyllenhali
2342 Pycnopterna striata
2339 Homodemus M-flavum
2281 Calocoris ochromelas
4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
331-332 Capsodes
334 Acetropis gimmerthalii
335 Stenodema calcarata
336 Stenodema trispinosa
337 Stenodema holsata
338 Stenodema laevigata
340 Notostira elongata
341 Megaloceraea recticornis
349 Leptopterna dolabrata
350 Leptopterna ferrugata
353 Himacerus apterus
354 Aptus mirmicoides
355 Anaptus major
356 Stalia boops
357 Nabicula lineata
358 Nabicula limbata
359 Nabicula flavomarginata
372 Anthocoris gallarumulmi
384 Ortus niger
386 Orius majusculus
387 Orius minutus
389 Lyctocoris campestris
390-392 Xylocoris
395 Dufouriellus ater
402-404 Empicoris
407 Rhynocoris annulatus
417 Piesma maculatum
418 Piesma quadratum
420-425 Berytinus
427 Gampsocoris punctipes
429 Lygaeus equestris
431 Nysius helveticus
435 Ortholomus punctipennis
438 Kleidocerys resedae
439 Kleidocerys truncatulus ericae
440 Cymus aurescens
450 Tropidophlebia costalis
456 Drymus latus
468 Lamproplax picea
462 Eremocoris abietis
465 Gastrodes abietum
466 Gastrodes grossipes
482 Stygnocoris sabulosus
483 Pachybrachius fracticollis
484 Pachybrachius luridus
485 Aellopus atratus
487 Graptopeltus lynceus
490 Peritrechus lundi
492-493 Rhyparochromus
494 Raglius alboacuminatus
495 Xantochilus quadratus
496-500 Megalonotus
502 Aphanus rolandri
508 Pterotmetus staphiliniformis
513-514 Dicranocephalus
2480-2483 Lopus
2492 Acetropis gimmerthali
2495 Stenodema calcaratum
2496 Stenodema trispinosum
2507 Stenodema holsatum
2503 Stenodema laevigatum
2510 Notostira erratica
2512 Megaloceroea linearis
2524 Maris dolobratus
2525 Maris ferrugatus
2006 Nabis apterus
2007 Nabis lativentris
2010 Nabis major
2012 Nabis boops
2017 Nabis lineatus
2016 Nabis limbatus
2019 Nabis flavomarginatus
2072 Anthocoris gallarum-ulmi
2090 Triphleps nigra
2095 Triphleps majuscula
2096 Triphleps minuta
2105 Lyctocorus campestris
2110-2118 Piezostethus
2135 Xylocoris ater
1738-1744 Ploiariola
1933 Rhinocoris tracundus
1423 Piesma maculata
1424 Piesma quadrata
1393-1409 Berytus
1421 Metacanthus punctipes
874 Spilostethus equestris
920 Nysius lineatus
925 Nysius punctipennis
937 Ischnorbynchus resedae
938 Ischnorbynchus ericae
934 Cymus obliquus
1038 Camptotelus lineolatus
1337 Drymus confusus
1346 Drymus piceus
1352 Eremocoris erraticus
1386 Gastrodes abietis
1387 Gastrodes ferrugineus
1199 Stygnocoris pedestris
1075 Pamera fracticollis
1078 Pamera lurida
1232 Microtoma atrata
1257 Aphanus lynceus
1214 Peritrechus sylvestris
1279-1280 Aphanus
1276 Aphanus alboacuminatus
1266 Aphanus quadratus
1099-1113 Rhyparochromus
1245 Calyptonotus rolandri
1139 Pterotmetus staphylinoides
765-768 Stenocephalus
AUKEMA:
517 Enoplops scapha
518 Coreus marginatus
519 Syromastus rhombeus
520 Spathocera dahlmannii
521 Arenocoris fallenu
522 Arenocoris waltlii
528 Corizus hyoscyami
529-531 Rhopalus
536 Legnotus limbosus
537 Legnotus picipes
542 Aethus flavicornis
544 Cydnus aterrimus
547 Odontoscelis lineola
552 Graphosoma lineatum
558 Eysarcoris aeneus
559 Eysarcoris fabricii
561-562 Holcostethus
574 Eurydema ventralis
For all species the presence or absence in the 12
Dutch provinces (fig. 1) is given. Questionable
records are indicated by a question mark. For rare,
new or endangered species (the species marked
with an asterisk (*) before their number) distribu-
tional data are treated in more detail under Re-
marks. In that section details on relevant literature,
synonymy, published records, unpublished records
and present status are given. Furthermore, distri-
bution maps (figs. 2-120) are given for most spe-
cies treated here (open circles: recorded before
1930; small dots: recorded between 1930 and 1960;
large dots: recorded since 1960; question marks:
doubtful records). Only records verified by the aut-
hor are included. Unless otherwise stated, the ma-
terial mentioned is deposited in the collection of
the collector. Collections are indicated by their ab-
breviations.
Material of the following Dutch collections is
included:
Museums and institutes: Zoologisch Museum,
Amsterdam (ZMA); Rijksmuseum van Natuur-
lijke Historie, Leiden (RML); Vakgroep Entomo-
logie, Landbouwuniversiteit, Wageningen (LUW);
Fries Natuurhistorisch Museum, Leeuwarden
(FNM); Natuurhistorisch Museum, Maastricht
(NMM); Rijksinstituut voor Natuurbeheer, Arn-
hem (RINA); Biologisch Station, Wijster (BSW);
Afdeling Bestrijding van Dierplagen, Ministerie
van VROM, Wageningen (ABW); Plantenziekten-
kundige Dienst Wageningen (PDW); Delta Insti-
tuut, Yerseke (DI).
Private collections: B. van Aartsen, 't Harde
(AA); B. Aukema, Wageningen (AU); J. Beernink,
Zaandam (BE); K. den Bieman, Bennekom (BI); R.
Checklist Hemiptera-Heteroptera
703 Coreus scapha
708 Syromastus marginatus
696 Verlusia rhombea
725 Spathocera dalmani
735 Pseudophloeus falleni
737 Pseudophloeus waltli
806 Therapha hyosciami
812-815 Corizus
46 Gnatoconus albomarginatus
47 Gnatoconus picipes
20 Cydnus flavicorne
39 Brachypelta aterrimus
112 Odontoscelis dorsalis
255 Graphosoma italicum
383 Eusarcoris aeneus
385 Eusarcoris melanocephalus
416-419 Peribalis
492 Eurydema ornatum
416. Aneurus laevis. Scale: 1 mm.
6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
H. Cobben (CB); C. Dolleman, Arnhem (D), Th.
Heijerman, Wageningen (HE), D. J. Hermes, Gel-
drop (HS); R. Ph. Jansen, Amsterdam (JA); G.
Morseld, Rheden (M); N. Nieser, Tiel (N); P. Poot,
Maastricht (P); H. Vallenduuk, Lelystad (VA); J. H.
Woudstra, Zaandam (W).
J. G. M. Cuppen (Vakgroep Hydrobiologie,
Landbouwuniversiteit, Wageningen), H. Moster-
dijk (Technologische Dienst Zeeuwse Waterschap-
pen (ZW), Terneuzen) and N. Nieser, Tilburg,
supplied many data for the Nepomorpha and Ger-
romorpha sections and the latter allowed the use of
data from the files of the Dutch section of the
European Invertebrate Survey, Leiden (marked
'EIS in the Remarks” section).
The reference section gives a complete list of
works containing records on Dutch Heteroptera
published since 1950. Publications not referred to
in the text are marked with an asterisk.
Fig. 1. Division of The Netherlands in 12 provinces. FR: Friesland; GR: Groningen; DR: Drenthe; OV: Overijssel;
FL: Flevoland; GL: Gelderland; UT: Utrecht; NH: Noord-Holland; ZH: Zuid-Holland; ZE: Zeeland; NB: Noord-
Brabant; LB: Limburg.
AUKEMA:
DIPSOCOROMORPHA
DIPSOCOROIDEA
Dipsocoridae
1
“2
Ceratocombus Signoret
coleoptratus (Zetterstedt)
Pachycoleus Fieber
waltli Fieber
NEPOMORPHA
NEPOIDEA
Nepidae
3
4
Nepinae
Nepa Linnaeus
cinerea Linnaeus
Ranatrinae
Ranatra Fabricius
linearis (Linnaeus)
CORIXOIDEA
Corixidae
#5
*6
10
Micronectinae
Micronecta Kirkaldy
Checklist Hemiptera-Heteroptera
CHECKLIST
FR GR DR OV FL GL UT NH ZH
subgenus Dichaetonecta Hutchinson
scholtzi (Fieber)
subgenus Micronecta s.s.
minutissima (Linnaeus)
poweri (Douglas & Scott)
Cymatiainae
Cymatia Flor
bonsdorffii (Sahlberg)
coleoptrata (Fabricius)
Corixinae
Glaenocorisa Thomson
propinqua propinqua (Fieber)
ZE
NB LB
eke
“pe
e e
e e
CE
CE
e e
CE
CE
8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
FR GR DR OV FL GL UT NH ZH ZE NB LB
Arctocorisa Wallengren
11 germari (Fieber) © - | 6... Orr Orr er Cy
Callicorixa White
12 praeusta praeusta (Fieber) 0... © ©. 6 © © ©) 50 Gener ine
Corixa Geoffroy
13 affinis Leach
14 dentipes (Thomson)
15 panzeri (Fieber)
16 punctata (Illiger)
Hesperocorixa Kirkaldy
17 castanea (Thomson)
18 linnaei (Fieber)
19 moesta (Fieber)
20 sahlbergi (Fieber)
Paracorixa Poisson
21 concinna concinna (Fieber) e: © © © © © © © nemen
Sigara Fabricius
subgenus Halicorixa Walton
*22 selecta (Fieber) u... -% Mr = = = NON
23 stagnalis stagnalis (Leach) e = {= =) 68 =. = eee
subgenus Microsigara Poisson
*24 hellensii (Sahlberg) EE — O ©
subgenus Pseudovermicorixa Jac-
zewski
25 nigrolineata nigrolineata (Fieber)
subgenus Retrocorixa Walton
26 limitata limitata (Fieber) e - © o e e oe e _ + o ©
27 semistriata (Fieber) eo o è è e. © © © © re hemme
subgenus Sigara s.s.
28 striata (Linnaeus)
subgenus Subsigara Stichel
29 distincta (Fieber)
30 falleni (Fieber)
31 fossarum (Leach) -
*32 jactans Jansson =e
*33 longipalis (Sahlberg) 1e ©, = | (ote) SMER
34 scotti (Fieber) o °.
subgenus Vermicorixa Walton
35 lateralis (Leach) e e oe ec eee © oe e © ©
NAUCOROIDEA
Naucoridae
Naucorinae
Ilyocoris Stàl
36 cimicoides (Linnaeus) o e o e o o o o e e © ©
AUKEMA: Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE NB IB
Naucoris Geoffroy
*37 maculatus Fabricius Se oes 2 © Sa) desen =
Aphelocheiridae
Aphelocheirus Westwood
*38 aestivalis (Fabricius) Zi i 0 are Ole
NOTONECTOIDEA
Notonectidae
Notonectinae
Notonecta Linnaeus
39 glauca glauca Linnaeus
40 lutea Miller
41 maculata Fabricius
42 viridis Delcourt
43 obliqua Thunberg
*44 reuteri Hungerford
PLEOIDEA
Pleidae
Plea Leach
45 minutissima Leach e eee © © e e © e oe e
GERROMORPHA
MESOVELIOIDEA
Mesoveliidae
Mesovelia Mulsant & Rey
46 furcata Mulsant & Rey © =. © © ce) “ee OOK eN ©
HEBROIDEA
Hebridae
Hebrus Curtis
subgenus Hebrus s.s.
47 pusillus (Fallen) e e o e e e ee © e oe ©
subgenus Hebrusella Poisson
48 ruficeps (Thomson) e + oe © e © e © © e ee @
HYDROMETROIDEA
Hydrometridae
10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
ERSGRSDREOM ELA GEMUTeNENZENZENNBMER
Hydrometra Latreille
49 gracilenta Horvth =. @ +6 © e @ @ ete Elie
50 stagnorum (Linnaeus) o e e e e e © oe © © ee ©
GERROIDEA
Veliidae
Veliinae
Velia Latreille
subgenus Plesiovelia Tamanini
51 caprai Tamanini 0 -.8 © © è © © vernemen e
*52 saulii Tamanini Le (Me
Microveliinae
Microvelia Westwood
*53 buenoi Drake = EL = #0 = er WE 2 O ©
*54 pygmaea (Dufour) Ee OO
55 reticulata (Burmeister) eo © © © © © © e © © © ©
Gerridae
Gerrinae
Gerris Fabricius
56 argentatus Schummel „eene 8 © è eneen je
57 gibbifer Schummel e — © +, = ©. e. ve) eg” neme
58 lacustris (Linnaeus) e e © © e © © © © e ©
*59 lateralis Schummel zee © ie ele
60 odontogaster (Zetterstedt) o © oe eo o e oe oe oe oe oe @
61 thoracicus Schummel e oe oe oe e o oe o o © @ ©
Aquarius Schellenberg
62 najas (De Geer) WW © + = © © © © Meo
63 paludum (Fabricius) e... (© è Korie Re
Limnoporus Stal
64 rufoscutellatus (Latreille) 0 «| © © = © eee 0...
LEPTOPODOMORPHA
LEPTOPODOIDEA
Saldidae
Chiloxanthinae
Chiloxanthus Reuter
65 pilosus (Fallén) © © vanen eee eee Noos
AUKEMA: Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE NB LB
Saldinae
Saldini
Salda Fabricius
66 littoralis (Linnaeus) en, ©, 00. ©’ Biu ee) =
*67 morio (Zetterstedt) ie
*68 muelleri (Gmelin) Oe A TA RA TA
Chartoscirtini
Halosalda Reuter
69 lateralis (Fallen) © © = =. 6. = puis gr0i sco) —
Saldula Van Duzee
subgenus Macrosaldula Southwood & Leston
#70 scotica (Curtis) SF Sete Se SOS ee er SOTO
subgenus Saldula s.s.
*7] arenicola (Scholtz) Oe
*72 c-album (Fieber) Ca Li
*73 fucicola (Sahlberg)
*74 melanoscela (Fieber)
*75 opacula (Zetterstedt)
76 orthochila (Fieber)
77 pallipes (Fabricius)
*78 palustris (Douglas)
79 pilosella (Thomson)
80 saltatorıa (Linnaeus)
e © e e © © © |
e e © | |
© e e © |
© © |
© o e © © |
e e ©
e e e e ee
e e e e © ©
© © e e © |
|
Micracanthia Reuter
*81 marginalis (Fallen) OT en OR)
Chartoscirta Stàl
82 cincta (Herrich-Schffer) e e oe oe oe © © © © © © ©
83 cocksii (Curtis) e - e o _
*84 elegantula (Fallen) ze Online
CIMICOMORPHA
TINGOIDEA
Tingidae
Tinginae
Campylosteira Fieber
85 verna (Fallen) © 0 let le
Acalypta Westwood
86 carinata (Panzer) =) linko: Weal (ee Cu is Konst lun 0
87 gracilis (Fieber) Re SS ee E
*88 marginata (Wolff) BE ar Ge E Or MO EN arena
*89 musci (Schrank) ENT ES ES RES e
12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
FR GR DR OV FL GL UT NH ZH ZE NB LB
#90 nigrina (Fallen) ze ee Dar wer UU = uL_
91 parvula (Fallen) o. 0 = + © “e “e Too
*92 platycheila (Fieber) o % Se & 0. ef, OR
Dictyonota Curtis
93 fuliginosa (Costa) =* -® 60% + : = © emmen hmnee
94 strichnocera (Fieber) Li ennn © eter kene
Kalama Puton
95 tricornis (Schrank) è. = = = = © + © rotaie) te
Derephysia Spinola
subgenus Derephysia s.s.
96 foliacea (Fallén) è. — © © = ‘e è. enmenmene
Galeatus Curtis
*97 maculatus (Herrich-Schäffer) Se SN Se ee SME
Stephanitis Stal
98 oberti (Kolenati) en OO
99 rhododendri Horváth Lu Et Lie e ale
Lasiacantha Stâl
*100 capucina (Germar) 2% 2806 Ee
Tingis Fabricius
subgenus Tropidocheila Fieber
*101 reticulata (Herrich-Schäffer) =sheLet = © inferi mens
subgenus Neolasiotropis Wagner
*102 pilosa Hummel = & & & © |, “Aa eee ©
subgenus Tingis s.s.
103 ampliata (Herrich-Schäffer) e: ‘o ‘è ‘è © © ‘è e. =e vee
104 cardui (Linnaeus) o © e © o © © e e © oe e@
*105 crispata (Herrich-Schäffer) Ee eee
Catoplatus Spinola
106 fabricii (Stal) & ea. e fe; eee Te re
Physatocheila Fieber
*107 costata (Fabricius) LL. £8 Ee
*108 dumetorum (Herrich-Schäffer) ee I OO
*109 smreczynskii (China) ee @ O one
Oncochila Stal
*110 simplex (Herrich-Schaffer) ai ta. ne ee > — ©
Dictyla Stal
111 convergens (Herrich-Schäffer) è = © © = © © © ‘o’ oe nce
112 echii (Schrank) EO Ot 0, TO
113 humuli (Fabricius) een. ans: 0 een
Agramma Stephens
114 laetum (Fallen) o 2.0. = © © ‘e;i’e;ize zone
AUKEMA: Checklist Hemiptera-Heteroptera
FReGRI DR OV FL GE Ul NH ZE ZE NB) EB
MIROIDEA
Microphysidae
Loricula Curtis
115 bipunctata (Perris) ores ce KO = Je) er eme menne
116 elegantula (Bärensprung)
117 pselaphiformis Curtis © = © © — © “e; olmo re; eo
Myrmedobia Bärensprung
118 coleoptrata (Fallén) © Ze NO Tete; VO ON vei ee
*119 distinguenda Reuter = — - —
120 exilis (Fallen) nee ae Leraren hide
©
I
|
|
|
I
I
Miridae
Bryocorinae
Bryocorini
Monalocoris Dahlbom
121 filicis (Linnaeus) o e oe o - © e e e e oe ©
Bryocoris Fallén
122 pteridis (Fallen) u. ae ©
Deraeocorinae
Clivinematini
Bothynotus Fieber
*123 pilosus (Boheman) EO)
Deraeocorini
Deraeocoris Kirschbaum
subgenus Camptobrochis Fieber
124 lutescens (Schilling) Os LO ee OO OT o 0 0
#125 punctulatus (Fallén) - - - = = ®
subgenus Deraeocoris s.s.
*126 annulipes (Herrich-Schäffer) ot =, 0 ee
127 cordiger (Hahn) HU I
*128 flavilinea (Costa) - = =
129 olivaceus (Fabricius) - = =
130 ruber (Linnaeus) oe t=
*131 scutellaris (Fabricius) - - +
132 trifasciatus (Linnaeus) - - 6
Alloeotomus Fieber
133 germanicus Wagner enter oe) oo) rene
134 gothicus (Fallén) e = © & = © © en è annie o
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Phylinae
Phylini
Lopus Hahn
decolor (Fallén)
Oncotylus Fieber
punctipes Reuter
viridiflavus (Goeze)
Conostethus Fieber
griseus (Douglas & Scott)
roseus (Fallén)
venustus (Fieber)
Hoplomachus Fieber
thunbergi (Fallén)
Megalocoleus Reuter
2 molliculus (Fallén)
3 pilosus (Schrank)
Amblytylus Fieber
albidus (Hahn)
brevicollis Fieber
nasutus (Kirschbaum)
Macrotylus Fieber
paykullii (Fallén)
solitarius (Meyer-Dür)
Orthonotus Stephens
rufifrons (Fallén)
Harpocera Curtis
thoracica (Fallén)
Tytthus Fieber
geminus (Flor)
pygmaeus (Zetterstedt)
Brachyarthrum Fieber
limitatum Fieber
Phylus Hahn
coryli (Linnaeus)
melanocephalus (Linnaeus)
palliceps (Fieber)
Plesiodema Reuter
pinetella (Zetterstedt)
FR GR DR OV FL GL UT NH ZH ZE
NB LB
o e
- e@
o) =
- ®@
Zale
e e
o e
o e
eo e
Te
SIRO
e
le
eo e
she
o e
e eo
er =
o oe
158
159
160
*161
162
165
164
165
166
167,
168
169
170
171
117?
1175)
174
175
176
La
178
179
*180
181
*182
2183
184
185
*186
187
188
AUKEMA:
Psallus Fieber
subgenus Apocremnus Fieber
betuleti (Fallén)
subgenus Mesopsallus Wagner
ambiguus (Fallén)
subgenus Phylidea Reuter
quercus (Kirschbaum)
subgenus Hylopsallus Wagner
assimilis Stichel
perrisi (Mulsant & Rey)
pseudoplatani Reichling
variabilis (Fallén)
wagneri Ossiannilsson
subgenus Psallus s.s.
albicinctus (Kirschbaum)
confusus Rieger
falleni Reuter
flavellus Stichel
haematodes (Gmelin)
lepidus Fieber
mollis (Mulsant & Rey)
salicis (Kirschbaum)
varians (Herrich-Schäffer)
subgenus Pityopsallus Wagner
luridus Reuter
Compsidolon Reuter
subgenus Coniortodes Wagner
salicellus (Herrich-Schäffer)
Phoenicocoris Reuter
obscurellus (Fallén)
Atractotomus Fieber
magnicornis (Fallén)
mali (Meyer-Dür)
parvulus Reuter
Plagiognathus Fieber
subgenus Poliopterus Wagner
albipennis (Fallén)
litoralis Wagner
subgenus Plagiognathus s.s.
alpinus (Reuter)
arbustorum (Fabricius)
chrysanthemi (Wolff)
fulvipennis (Kirschbaum)
subgenus Parapsallus Wagner
vitellinus (Scholtz)
Chlamydatus Curtis
subgenus Euattus Kerzhner
pulicarius (Fallén)
Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE NB LB
16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
FR GR DR OV FL GL UT NH ZH ZE NB IB
189 pullus (Reuter) a8 ei, ©. Degree
subgenus Chlamydatus s.s.
190 saltitans (Fallen) e - e e ee è o oe oo oe @
subgenus Eurymerocoris Kirsch-
baum
*191 evanescens (Boheman) BEE NN DO. OOO
Monosynamma Scott
*192 bohemanni (Fallen) ei = .© == © = eten enmenme
193 maritimum Wagner © = dre + (58 ennen
*194 sabulicola Wagner = = + = © © Vier
Campylomma Reuter
*195 annulicorne (Signoret) a= 6 2 © © © 0
196 verbasci (Meyer-Dür) zé © + - © „elit MO
Criocoris Fieber
*197 crassicornis (Hahn) e > Zie
*198 sulcicornis (Kirschbaum) pe We de
Salicarus Kerzhner
199 roseri (Herrich-Schäffer) 2 © = ne © © 6,0.) 0 tete e
Sthenarus Fieber
200 rotermundi (Scholtz) 0 slee © © © © Mere “ae
Asciodema Reuter
*201 obsoletum (Fieber) In 0 070 Oo
Hallodapini
Hallodapus Fieber
*202 rufescens (Burmeister) = = © © =| er a = —
Systellonotus Fieber
203 triguttatus (Linnaeus) EEE OO
Cremnocephalus Fieber
*204 albolineatus Reuter sn © _ 0 =
Dicyphinae
Macrolophus Fieber
205 nubilus (Herrich-Schäffer) LE et © Lele @ 0
Dicyphus Fieber
subgenus Dicyphus s.s.
*206 constrictus (Boheman) 2 = en wl dE
207 epilobii Reuter O = 0 e: e è je, enge
208 errans (Wolff) so 2 ie var © té eee ee
209 pallidus (Herrich-Schäffer) ROO hole
subgenus Idolocoris Douglas & Scott
OS ne
210 pallicornis (Meyer-Dür) © %-. © Je =" ee
211
212
2115)
228
*229
230
AUKEMA:
subgenus Brachycerea Fieber
annulatus (Wolff)
globulifer (Fallén)
Campyloneura Fieber
virgula (Herrich-Schäffer)
Orthotylinae
Pilophorini
Hypseloecus Reuter
visci (Puton)
Pilophorus Hahn
cinnamopterus (Kirschbaum)
clavatus (Linnaeus)
confusus (Kirschbaum)
perplexus (Douglas & Scott)
Halticocorini
Halticus Hahn
apterus (Linnaeus)
luteicollis (Panzer)
saltator (Geoffroy)
Strongylocoris Blanchard
leucocephalus (Linnaeus)
luridus (Fallén)
Pachytomella Reuter
parallela (Meyer-Diir)
Orthocephalus Fieber
coriaceus (Fabricius)
saltator (Hahn)
Orthotylini
Reuteria Puton
marqueti Puton
Malacocoris Fieber
chlorizans (Panzer)
Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE NB LB
Fieberocapsus Carvalho & Southwood
flaveolus (Reuter)
Cyllecoris Hahn
histrionicus (Linnaeus)
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Dryophilocoris Reuter
flavoquadrimaculatus (De Geer)
Globiceps Le Peletier & Serville
subgenus Paraglobiceps Wagner
2 flavomaculatus (Fabricius)
fulvicollis cruciatus Reuter
subgenus Globiceps s.s.
sphegiformis (Rossi)
Heterocordylus Fieber
genistae (Scopoli)
leptocerus (Kirschbaum)
tibialis (Hahn)
tumidicornis (Herrich-Schäffer)
Heterotoma Le Peletier & Serville
planicornis (Pallas)
Blepharidopterus Kolenati
angulatus (Fallén)
diaphanus (Kirschbaum)
Orthotylus Fieber
subgenus Pinocapsus Southwood
fuscescens (Kirschbaum)
subgenus Orthotylus s.s.
flavinervis (Kirschbaum)
marginalis Reuter
nassatus (Fabricius)
prasinus (Fallén)
tenellus (Fallén)
virens (Fallén)
viridinervis (Kirschbaum)
subgenus Litocoris Fieber
ericetorum (Fallén)
subgenus Neopachylops Wagner
adenocarpi (Perris)
concolor (Kirschbaum)
virescens (Douglas & Scott)
subgenus Melanotrichus Reuter
flavosparsus (Sahlberg)
moncreaffi (Douglas & Scott)
rubidus (Fieber)
Pseudoloxops Kirkaldy
coccineus (Meyer-Diir)
Cyrtorhinus Fieber
caricis (Fallén)
Neomecomma Southwood
bilineatum (Fallén)
FR GR DR OV FL GL UT NH ZH ZE
© e © |
NB LB
© ® ® © ® e ©
e © © = = e ®
© ® ® © e © ®
®
= = e = = = ©
© ©
© © ® © ® © e
= = = = ©
© © © ® © ® ©
© © e © ® e ®
® © © ® © = ®
© = = = = = ©
© © © ® © = ®
© © © © © © ©
© © ® © ® = ©
© = © ® = = ©
© © ® © = = ©
à = = = = = ©
© © © © © = ®
© © © = = ® ®
© © e = = ©
© © © © = © ®
® © © © = e e
® © © ® ® = ©
= = © = ® = =
= = e = e =
e © e ® © = ©
© = = = = = =
® = e = = ® ©
*260
*261
262
AUKEMA:
Mecomma Fieber
subgenus Mecomma s.s.
ambulans (Fallén)
Mirinae
Pithanini
Myrmecoris Gorski
gracilis (Sahlberg)
Pithanus Fieber
maerkelii (Herrich-Schäffer)
Mirini
Lygus Hahn
adspersus (Schilling)
gemellatus (Herrich-Schäffer)
maritimus Wagner
pratensis (Linnaeus)
rugulipennis Poppius
wagneri Remane
Liocoris Fieber
tripustulatus (Fabricius)
Orthops Fieber
basalis (Costa)
campestris (Linnaeus)
kalmii (Linnaeus)
Pinalitus Kelton
atomarius (Meyer-Dür)
cervinus (Herrich-Schäffer)
rubricatus (Fallén)
viscicola (Puton)
Plesiocoris Fieber
minor Wagner
rugicollis (Fallén)
Lygocoris Reuter
subgenus Lygocoris s.s.
pabulinus (Linnaeus)
subgenus Neolygus Knight
contaminatus (Fallén)
populi Leston
viridis (Fallén)
subgenus Apolygus China
limbatus (Fallén)
lucorum (Meyer-Diir)
rhamnicola (Reuter)
spinolae (Meyer-Diir)
Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE
© © | © |
NB LB
Ere
o oe
e e
Zu |
e e
eo e
e e
e ©
e e
o e
o e
. e
oo.
.e e
=. ©
3 di
eo
e e
°
e e
eo e
- e@
SI
19
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Agnocoris Reuter
reclairei Wagner
rubicundus (Fallén)
Camptozygum Reuter
aequale (Villers)
Polymerus Hahn
subgenus Poeciloscytus Fieber
palustris (Reuter)
unifasciatus (Fabricius)
2 vulneratus (Panzer)
subgenus Polymerus s.s.
holosericeus Hahn
nigrita (Fallén)
Charagochilus Fieber
gyllenhalii (Fallén)
Dichrooscytus Fieber
intermedius Reuter
rufipennis (Fallén)
Miris Fabricius
striatus (Linnaeus)
Hadrodemus Fieber
m-flavum (Goeze)
Calocoris Fieber
subgenus Trichocalocoris Wagner
pilicornis (Panzer)
schmidtii (Fieber)
subgenus Rhabdomiris Wagner
striatellus (Fabricius)
subgenus Closterotomus Fieber
biclavatus (Herrich-Schäffer)
fulvomaculatus (Villers)
subgenus Calocoris s.s.
affinis (Herrich-Schäffer)
norvegicus (Gmelin)
roseomaculatus (De Geer)
Adelphocoris Reuter
quadripunctatus (Fabricius)
lineolatus (Goeze)
seticornis (Fabricius)
ticinensis (Meyer-Diir)
Megacoelum Fieber
2 beckeri (Fieber)
infusum (Herrich-Schäffer)
FR GR DR OV FL GL UT NH ZH ZE NB LB
314
II
316
#17
318
319
320
321
322
323
*324
325
326
327
3928
329
330
Jo
2532
339
*334
358
*336
Do)
338
250)
340
*341
AUKEMA:
Stenotus Jakovlev
binotatus (Fabricius)
Miridius Fieber
quadrivirgatus (Costa)
Phytocoris Fallén
subgenus Phytocoris s.s.
dimidiatus Kirschbaum
intricatus Flor
longipennis Flor
pini Kirschbaum
populi (Linnaeus)
reuteri Saunders
tiliae (Fabricius)
subgenus Ktenocoris Wagner
insignis Reuter
singeri Wagner
ulmi (Linnaeus)
varipes (Boheman)
Capsus Fabricius
ater (Linnaeus)
pilifer Remane
wagneri Remane
Pantilius Curtis
tunicatus (Fabricius)
Capsodes Dahlbom
gothicus (Linnaeus)
sulcatus (Fieber)
Stenodemini
Acetropis Fieber
carinata (Herrich-Schäffer)
gimmerthalii (Flor)
Stenodema Laporte
subgenus Brachystira Fieber
calcarata (Fallén)
trispinosa Reuter
subgenus Stenodema s.s.
holsata (Fabricius)
laevigata (Linnaeus)
virens (Linnaeus)
Notostira Fieber
elongata (Geoffroy)
Megaloceroea Fieber
recticornis (Geoffroy)
Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE NB LB
21
22. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
FR GR DR OV FL GL UT NH ZH ZE NB LB
Trigonotylus Fieber
*342 caelestialium (Kirkaldy) eo © © eee eee e oe ©
343 psammaecolor Reuter EO Oes
344 pulchellus (Hahn) zie = © = © © Rene
*345 ruficornis (Geoffroy) o + o e oe e e e o è. e
Teratocoris Fieber
346 antennatus (Boheman) e = + = =. © Le renten mormel ec
*347 paludum Sahlberg ==]. 6@@2 © (eae
*348 saundersi Douglas & Scott SM Lie = = = > © Relea
Leptopterna Fieber
349 dolobrata (Linnaeus) en nette _ © (uao
350 ferrugata (Fallén) CC 0. o e o ee e e e
Isometopinae
Isometopus Fieber
*35] intrusus (Herrich-Schäffer) ziet = © + meme
CIMICOIDEA
Nabidae
Prostemmatinae
Prostemmatini
Prostemma Laporte
*352 guttula (Fabricius) D © = ©
Nabinae
*Nabini
Himacerus Wolff
353 apterus (Fabricius) o e © e ei eier ©
Aptus Hahn
354 mirmicoides (Costa) e e © © = è è. 0e 50. eer ©
Anaptus Kerzhner
355 major (Costa) 9 (e (00 8 © e eetmenmen e
Stalia Reuter
*356 boops (Schiödte) evet - © © „erelkkemieme
Nabicula Kirby
subgenus Limnonabis Kerzhner
357 lineatus (Dahlbom) o © “e te © © © veren emmeme
subgenus Dolichonabis Reuter
358 limbatus (Dahlbom) e ee ee © © eee © e
subgenus Nabicula s.s.
359 flavomarginatus (Scholtz) o e o oe _ e © eo o © oo @
AUKEMA:
Nabis Latreille
brevis Scholtz
ericetorum Scholtz
ferus (Linnaeus)
pseudoferus Remane
rugosus (Linnaeus)
Anthocoridae
365
*366
367
*368
#369
2370
DI
372
215)
*374
DII
376
a
378
a)
*380
381
*382
583
384
*385
386
387
*388
Anthocorinae
Anthocorini
Temnostethus Fieber
gracilis (Horvath)
longirostris (Horvath)
pusillus (Herrich-Schäffer)
Elatophilus Reuter
nigricornis (Zetterstedt)
Anthocoris Fallén
amplicollis Horváth
butleri Le Quesne
confusus Reuter
gallarumulmi De Geer
limbatus Fieber
minki minki Dohrn
nemoralis (Fabricius)
nemorum (Linnaeus)
pilosus (Jakovlev)
sarothamni Douglas & Scott
simulans Reuter
visci Douglas
Tetraphleps Fieber
bicuspis (Herrich-Schäffer)
Acompocoris Reuter
alpinus Reuter
pygmaeus (Fallén)
Oriini
Orius Wolff
subgenus Orius s.s.
niger (Wolff)
subgenus Heterorius Wagner
laticollis (Reuter)
majusculus (Reuter)
minutus (Linnaeus)
vicinus (Ribaut)
Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE NB LB
© © |
23
389
390
391
992
>95
3094
395
06
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Lyctocorinae
Lyctocorini
Lyctocoris Hahn
campestris (Fabricius)
Xylocorini
Xylocoris Dufour
subgenus Proxylocoris Carayon
galactinus (Fieber)
subgenus Xylocoris s.s.
cursitans (Fallén)
formicetorum (Boheman)
Cardiastethini
Brachysteles Mulsant & Rey
parvicornis (Costa)
Cardiastethus Fieber
fasciiventris (Garbiglietti)
Dufouriellus Kirkaldy
ater (Dufour)
Scolopini
Scoloposcelis Fieber
pulchella (Zetterstedt)
Cimicidae
>97
398
5299)
*400
*401
Oeciacus Stâl
hirundinis (Lamarck)
Cimex Linnaeus
lectularius Linnaeus
columbarius Jenyns
dissimilis (Horváth)
pipistrelli Jenyns
REDUVOIDEA
Reduviidae
*402
403
404
Emesinae
Empicoris Wolff
baerensprungi (Dohrn)
culiciformis (De Geer)
vagabundus (Linnaeus)
FR GR DR OV FL GL UT NH ZH ZE
NB LB
e
ee
o e
©
sure
oo
o e
o ©
°
oo
*405
406
*407
*408
*409
*410
AUKEMA:
Stenopodainae
Pygolampis Germar
bidentata (Goeze)
Reduviinae
Reduvius Fabricius
personatus (Linnaeus)
Harpactorinae
Rhynocoris Hahn
annulatus (Linnaeus)
Coranus Curtis
subapterus (De Geer)
woodroffei Putshkov
Phymatinae
Phymata Latreille
crassipes (Fabricius)
PENTATOMORPHA
ARADOIDEA
Aradidae
411
*412
413
*414
*415
*416
Aradinae
Aradus Fabricius
cinnamomeus (Panzer)
corticalis (Linnaeus)
depressus (Fabricius)
signaticornis Sahlberg
Aneurinae
Aneurus Curtis
avenius (Dufour)
laevis (Fabricius)
PIESMATOIDEA
Piesmatidae
417
*418
*419
Piesma Le Peletier & Serville
maculatum (Laporte)
quadratum (Fieber)
salsolae (Becker)
Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE NB LB
26
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
COROIDEA
Berytidae
420
421
422
423
*424
425
426
427
*428
Berytinae
Berytinus Kirkaldy
subgenus Berytinus s.s.
clavipes (Fabricius)
hirticornis (Brullé)
minor (Herrich-Schäffer)
subgenus Lizinus Mulsant & Rey
crassipes (Herrich-Schäffer)
montivagus (Meyer-Dür)
signoreti (Fieber)
Neides Latreille
tipularius (Linnaeus)
Metacanthinae
Gampsocoris Fuss
punctipes (Germar)
Metatropis Fieber
rufescens (Herrich-Schäffer)
Lygaeidae
*429
*430
Lygaeinae
Lygaeus Fabricius
equestris (Linnaeus)
Spilostethus Stäl
saxatilis Scopoli
Orsillinae
Nysius Dallas
subgenus Macroparius Stàl
helveticus (Herrich-Schäffer)
subgenus Nysius s.s.
ericae (Schilling)
thymi (Wolff)
subgenus Tropinysius Wagner
senecionis (Schilling)
Ortholomus Stàl
punctipennis (Herrich-Schäffer)
Orsillus Dallas
depressus Dallas
FR GR DR OV FL GL UT NH ZH ZE NB LB
= = = = = © = = = ® = ©
= = = = = © = = = © e
© = ® ® = © © © ® © ®
= = © = = ® © © © ® e ©
= = = = = = = = = ® > =
© = © = = © © ® © ® ©
© = = = = © ® ® © © © ©
® = = = = = = ® © ® = ©
= = = © = = e
? = = = = = = = = = = ®
= = = = = = = = = = = ©
© © © © = © © © = = e ©
© = © © = © © = = ® ®
© ® © © © ® e © ® © ©
© ® © = = ® e © ® © e ©
*437
438
#459
*440
441
442
*443
444
*445
446
*447
*448
*449
*450
451
452
AUKEMA:
Ischnorhynchinae
Kleidocerys Stephens
privignis Horvath
resedae (Panzer)
truncatulus ericae (Horvath)
Cyminae
Cymus Hahn
aurescens Distant
claviculus (Fallén)
glandicolor Hahn
melanocephalus Fieber
Blissinae
Ischnodemus Fieber
sabuleti (Fallén)
Geocorinae
Geocoris Fallén
ater (Fabricius)
grylloides (Linnaeus)
megacephalus (Rossi)
Oxycareninae
Metopoplax Fieber
ditomoides (Costa)
Oxycarenus Fieber
modestus (Fallén)
Tropidophlebia Kerzhner
costalis (Herrich-Schäffer)
Artheneinae
Chilacis Fieber
typhae (Perris)
Heterogastrinae
Heterogaster Schilling
urticae (Fabricius)
Rhyparochrominae
Plinthisini
Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE NB LB
27
28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
FR GR DR OV FL GL UT NH ZH ZE NB IB
Plinthisus Stephens
453 brevipennis (Latreille) è = © © = è © + el etre 1e
*454 pusillus (Scholtz) RC. LO =
Antillocorini
Tropistethus Fieber
*455 holosericeus (Scholtz) Et ee N = ©
Drymini
Drymus Fieber
subgenus Drymus s.s.
*456 latus Douglas & Scott Mena nn = ede
*457 pilicornis (Mulsant) leo e «= sel. 8S S00 See
subgenus Sylvadrymus Le Quesne
458 brunneus (Sahlberg) ee + © è + © © ‘olio mono
*459 pumilio Puton = = == enfin *e
*460 ryeii Douglas & Scott o e oe e oe e o o e e oe ©
461 sylvaticus (Fabricius) e - © e e o © eo o © e ©
Eremocoris Fieber
462 abietis (Linnaeus) e. = = = = © “©, Se nhemtenmen te
463 plebejus (Fallen) è. + — © è © {feet} tes ©
*464 podagricus (Fabricius) + OO = ©
Gastrodes Westwood
465 abietum Bergroth e - @© - - + © © zuende
466 grossipes (De Geer) e - © e © o o o e oo e ©
Ischnocoris Fieber
467 angustulus (Boheman) CCC OS, — OO
Lamproplax Douglas & Scott
*468 picea (Flor) © 2. & = + et = _ O
Scolopostethus Fieber
469 affinis (Schilling) e e © oe e e o © oe © ee ©
470 decoratus (Hahn) ee = © © © © © © “el ele
471 pictus (Schilling) Eee o + ee © oo
*472 pilosus (Reuter) Ber VE ve ARIES”
*473 pseudograndis Wagner Eee
474 puberulus Horvath = = =< ee (ee
475 thomsoni Reuter e + e © e e o © © © oe @
Taphropeltus Stal
476 contractus (Herrich-Schäffer) L Mi. Me 4m = “è. © Toniolo nmionine
*477 hamulatus (Thomson) - © - @
Stygnocorini
Acompus Fieber
478 rufipes (Wolff) = © © © = © ‘ee (eee
AUKEMA: Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE NB LB
Lasiosomus Fieber
*479 enervis (Herrich-Schäffer) Se ee
Stygnocoris Douglas & Scott
480 fuligineus (Geoffroy) 0. = -e: © è. © ee e we ©
481 rusticus (Fallén) è =. 2 +, © de! “eee 'erthel.:e
482 sabulosus (Schilling) è: © © © =, © è enj eenen ie
Myodochini
Pachybrachius Hahn
483 fracticollis (Schilling) © = le © = © © eee
*484 luridus Hahn = Ta ie VOO ORNE ENEN =
Rhyparochrominae
Rhyparochromini
Aellopus Wolff
*485 atratus (Goeze) weise Ey ee PE a lea
Beosus Amyot & Serville
486 maritimus (Scopoli) o e + + + e e
Graptopeltus Stàl
487 lynceus (Fabricius) Oene er 0) ne) le Terre
Peritrechus Fieber
*488 angusticollis (Sahlberg)
489 geniculatus (Hahn)
*490 lundii (Gmelin) Coltan alia =
491 nubilus (Fallen) © = ve: =
Rhyparochromus Hahn
492 phoeniceus (Rossi) OH en SD LO. © One nie ©
493 pini (Linnaeus) è... Lo =" OO Kermi Mi mken 6
Raglius Stàl
*494 alboacuminatus (Goeze) SSS ie
Xanthochilus Stàl
495 quadratus (Fabricius) et CAM LS Te. © el ijt Mierde
Megalonotini
Megalonotus Fieber
*496 antennatus (Schilling) e ene. Te
497 chiragra (Fabricius) ®
498 dilatatus (Herrich-Schäffer) en
499 praetextatus (Herrich-Schäffer)
*500 sabulicola (Thomson)
®
I
I
|
|
© © © |
© |
© © |
Sphragisticus Stal
501 nebulosus (Fallen) e e o o oe © © e © © ©
30
502
*503
*504
505
506
507
508
509
2510
Si
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Gonianotini
Aphanus Laporte
rolandri (Linnaeus)
Emblethis Fieber
griseus (Wolff)
verbasci (Fabricius)
Gonianotus Fieber
marginepunctatus (Wolff)
Macrodema Fieber
microptera (Curtis)
Pionosomus Fieber
varius (Wolff)
Pterotmetus Fieber
staphiliniformis (Schilling)
Trapezonotus Fieber
arenarius (Linnaeus)
desertus Seidenstücker
dispar Stal
Pyrrhocoridae
UZ
Pyrrhocoris Fallén
apterus (Linnaeus)
Stenocephalidae
253013
*514
Dicranocephalus Hahn
agilis (Scopoli)
medius (Mulsant & Rey)
Coreidae
SD
SG
DIL]
518
519
Coreinae
Gonocerus Latreille
acuteangulatus (Goeze)
juniperi Herrich-Schäffer
Enoplops Amyot & Serville
scapha (Fabricius)
Coreus Fabricius
marginatus (Linnaeus)
Syromastus Berthold
rhombeus (Linnaeus)
FR GR DR OV FL GL UT NH ZH ZE
NB LB
o e
et =
.e oe
eo
eo
er
DIO
2,3
© e
eo eo
SRO)
= lo
eo e@
. e
e e
eo
SAI
2522
525)
AUKEMA:
Spathocera Stein
dahlmannii (Schilling)
Pseudophloinae
Arenocoris Hahn
fallenii (Schilling)
waltlii (Herrich-Schäffer)
Bathysolen Fieber
nubilus (Fallén)
Ceraleptus Costa
lividus Stein
Coriomeris Westwood
denticulatus (Scopoli)
Alydidae
526
Alydus Fabricius
calcaratus (Linnaeus)
Rhopalidae
Rhopalinae
Rhopalini
Liorhyssus Stal
hyalinus (Fabricius)
Corizus Fallén
hyoscyami (Linnaeus)
Rhopalus Schilling
subgenus Rhopalus s.s.
parumpunctatus Schilling
subrufus (Gmelin)
subgenus Aeschyntelus Stal
maculatus (Fieber)
Chorosomatini
Myrmus Hahn
miriformis (Fallén)
Chorosoma Curtis
schillingi (Schilling)
Checklist Hemiptera-Heteroptera
FR GR DR OV FL GL UT NH ZH ZE
NB LB
e e
pio,
e e
o e
o e
eo e
e e
= (®
. e
o oe
e e
>il
32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
FR GR DR OV FL GL UT NH ZH ZE NB LB
Maccevethini
Stictopleurus Stàl
*534 abutilon abutilon (Rossi) = 2 4 4 © Sirene
*535 punctatonervosus (Goeze) ze Æ =. © Ste
PENTATOMOIDEA
Cydnidae
Sehirinae
Legnotus Schioedte
536 limbosus (Geoffroy) 0. 5 592 32 © © ermee
537 picipes (Fallén) eo. = © = = © = ele ee
Sehirus Amyot & Serville
subgenus Tritomegas Amyot & Ser-
ville
538 bicolor (Linnaeus) © © © = © © ere mememe
subgenus Adomerus Mulsant & Rey
539 biguttatus (Linnaeus) Ss © © + + ee mie de
subgenus Sehirus s.s.
540 luctuosus Mulsant & Rey ® = = = = © © @ oz
#54] morio (Linnaeus) *. a £& oo & Se ae
Cydninae
Aethus Dallas
42 flavicornis (Fallen) EL O = =
3 nigrita (Fallén) ORO OO OO
Cydnus Fabricius
*544 aterrimus (Forster) = a ka (La ee Ro
Thyreocoridae
Thyreocoris Schrank
545 scarabaeoides (Linnaeus) OCR OO
Scutelleridae
Odontoscelis Laporte
546 fuliginosa (Linnaeus) TO O 10
#547 lineola (Rambur) eee Ye OO OO en
Phimodera Germar
*548 humeralis (Dalman) Oe
Eurygaster Laporte
#549 austriaca (Schrank) Sa ses ar ee emmen = 0
550 maura (Linnaeus) 2 ae = Me
551 testudinaria (Geoffroy) = 2 2 £ fa, © © ‘© Eee
AUKEMA:
Pentatomidae
552
553
554
III
556
DI
558
220)
*560
OI
#562
563
*564
509
#566
Don
568
569
570
Podopinae
Graphosoma Laporte
lineatum (Linnaeus)
Podops Laporte
inuncta (Fabricius)
Pentatominae
Sciocoris Fallén
cursitans (Fabricius)
Aelia Fabricius
acuminata (Linnaeus)
klugii Hahn
Neottiglossa Kirby
pusilla (Gmelin)
Eysarcoris Hahn
aeneus (Scopoli)
fabricii (Kirkaldy)
Rubiconia Dohrn
intermedia (Wolff)
Holcostethus Fieber
sphacelatus (Fabricius)
vernalis (Wolff)
Palomena Mulsant & Rey
prasina (Linnaeus)
viridissima (Poda)
Chlorochroa Stal
juniperina (Linnaeus)
pinicola (Mulsant & Rey)
Carpocoris Kolenati
fuscispinus (Boheman)
purpureipennis (De Geer)
Dolycoris Mulsant & Rey
baccarum (Linnaeus)
Piezodorus Fieber
lituratus (Fabricius)
Pentatoma Olivier
rufipes (Linnaeus)
Checklist Hemiptera-Heteroptera
FREGREDREOV FER GEZUNZNEEZHTZEINBETB
33
34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
FR GR DR OV FL GL UT NH ZH ZE NB LB
Eurydema Laporte
*572 dominulus (Scopoli) ee en en Rene
573 oleracea (Linnaeus) 0. = @ = è è (open
*574 ventralis (Kolenati) Spa & & & & Zen
Amyotinae
Picromerus Amyot & Serville
575 bidens (Linnaeus) o o oe e o e e oe e o o ©
Arma Hahn
576 custos (Fabricius) ZI = © © © © ee eee Lie
Troilus Stâl
577 luridus (Fabricius) è" — © © ‘+ © ‘© © eo. = we we
Rhacognathus Fieber
578 punctatus (Linnaeus) ers. è te = © 6. © emtheumene
Jalla Hahn
*579 dumosa (Linnaeus) ee Be en LO
Zicrona Amyot & Serville
580 caerulea (Linnaeus) e + e e © oe o e e _ e o.
Acanthosomatidae
Acanthosomatinae
Acanthosoma Curtis
581 haemorrhoidale (Linnaeus) ® . 0 © © © ©) selon e
Cyphostethus Fieber
582 tristriatus (Fabricius) è © + = emerson
Elasmostethus Fieber
583 interstinctus (Linnaeus) © © © ° © © © ©, Net Moie
Elasmuchinae
Elasmucha Stàl
584 ferrugata (Fabricius) Ren OO
585 fieberi (Jakovlev) e
586 grisea (Linnaeus) e e oe © e e © © o © © @
AUKEMA:
Fig. 2. Micronecta scholtzi. Symbols on this and following
maps: open circles: recorded before 1930 only; small dots:
recorded between 1930 and 1960; large dots: recorded
since 1960; ?: doubtful record.
Fig. 3. Micronecta minutissima.
Checklist Hemiptera-Heteroptera 35
REMARKS
2. Pachycoleus waltli
References. - Reclaire (1936: 256, as P. rufescens Sahl-
berg; 1940: 113).
Published records. - ® Limburg: Spaubeek (Reclaire
1936); Wijlre (Reclaire 1940).
Unpublished records. - ® Limburg: Noorbeek, iii-
half.iv.1960, 1 6,2 Q and 1 larva, Br. Arnoud; Ravensbos,
13.1x.1961, R. H. Cobben; Herkenbosch, Meijnweg,
16.ix.1961, R. H. Cobben; Elsloo, 11.v. 1967, P. J. Brak-
man (RML) & P. Poot.
5. Micronecta scholtzi (fig. 2)
Synonym: M. meridionalis Costa (Jansson 1986).
References. - Reclaire (1932: 235; 1940: 115, as mert-
dionalis Costa; 1943: 121); Nieser (1982: 61-62, as me-
ridionalis).
Published records. - ® Gelderland: Nijmegen (Nieser
1982). ® Utrecht: Abcoude, 13.vii.1936, G. Kruseman
(ZMA) (Reclaire 1943). ® Z-Holland: Leiden (Reclaire
1932); Warmond, Poelsmeer, D. C. Geijskes (Reclaire
1940). ® Zeeland: Kwadendamme, 28.vi.1977 (DI) (Nie-
ser 1982; !EIS); Nisse, 28.vi.1977 (DI) (Nieser 1982;
lEIS).
Unpublished records. - ® Overijssel: Zwolle, Schelle,
5.vi.1988, B. Aukema. ® Gelderland: Lobith, 18.v.1959
(K.W.R. Zwart); Wageningen, 11.ix.1971, R. H. Cobben;
Lienden, 28.vii.1981, K. den Bieman; Hemmen,
29.v.1985, B. Aukema; Wamel, 16.ix.19, K. den Bieman;
Winterswijk, Italiaanse Meren, 25.vii.1986, D. J. Hermes.
© Utrecht: Rhenen, 1.vi.1985, B. Aukema; Ibid., Blauwe
Kamer, 29.vii.1985, B. Aukema & J. H. Woudstra. ®
Zeeland: ‘s-Gravenpolder, 28.vii & 5.x.1976, DI, N.
Tramper; Nisse, 30.vii.1976, DI, N. Tramper; Ibid., 8.iii
& 28. vi.1977, DI, I. J. Weeber; Ibid., 27.vii & 11.x.1978,
DI, S. van der Boog; Zuiddorpe, 2.vii.1981, DI, B. Krebs;
Rilland, 29.ix.1988, ZW, H. Mosterdijk; Aardenburg, 1
locality, Axel, 7 localities, Hontenisse, 1 locality, Hulst, 5
localities and Oostburg, 2 localities, 1986-1988, ZW, H.
Mosterdijk; © N-Brabant: Drunen, Zeedijk, 28. vii. 1981,
Nieser et al. (!EIS). © Limburg: Echt, 27.viii.1975, E. J.
van Nieukerken (RML).
6. Micronecta minutissima (fig. 3)
References. - Reclaire (1932: 235); Nieser (1982: 62).
Published records. - ® Overijssel: Ommen, vi.1916, D.
MacGillavry (ZMA) (Reclaire 1932; 1943, as M. powers).
© Gelderland: Buren (Nieser 1982); ® Utrecht: Maarse-
veen (Nieser 1982). ® Z-Holland: Voorne (Nieser 1982);
© Zeeland: 's Gravenpolder, Nisse (Nieser 1982; !EIS).
© N-Brabant: Deurne, 26.v.1920, G. Romijn (ZMA) (Re-
claire 1932).
Unpublished records. - ® Groningen: Haren,
S.vi.1976, Beerta, 8.vi.1988, Winschoten, 8.vi.1988, B.
Aukema. ® Drenthe: Zuidlaren, vii.1956, A. J. Besseling
(K. W. R. Zwart, RML, ZMA, P). ® Overijssel: Zwolle,
Schelle, 5.vi.1988, B. Aukema. ® Gelderland: Lobith,
1959.v.18 (K. W. R. Zwart); Lienden, 3.vii.1972, R. H.
Cobben; Ubbergen, Groenlanden, 1.vi.1974, E. J. van
Nieukerken (RML); Echteld, Ochten, 8.vii.1975, B. Drost
36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
(N). ® Z-Holland: Tienhoven, 21.v.1981, N. Nieser et al.
(!EIS). ® N-Brabant: Drunen, Zeedijk, 28.vii.1981, N.
Nieser et al. (!EIS). ® Limburg: Roermond, 6.viii.1946,
R. H. Cobben (Reclaire 1951, as M. powers).
7. Micronecta poweri
References. — Reclaire (1943: 121); Nieser (1982: 62-
63).
Published records. - © Overijssel: Oud-Ootmarsum,
28.v.1966, N. Nieser (Nieser 1982; !EIS). © Gelderland:
Winterswijk, 15-21.vi.1921, 1 4, 1 9, P. van der Wiel
(ZMA) (Reclaire 1943). © N-Brabant: Oirschot, vii.1908,
D. MacGillavry (ZMA) (Reclaire 1932, as M. minutis-
sima; 1943).
Present status. — Very rare, not recorded since 1966.
22. Sigara selecta (fig. 4)
References. - Fokker (1891: 360); Duffels (1962); Hi-
gler & Duffels (1965: 109); Krebs (1982).
Note. - The records from Hilversum, Bergen op Zoom
and Halsteren in Reclaire (1932: 229) concern other
species (Reclaire 1943; Duffels 1962).
Published records. — ® Friesland: Terschelling (Duffels
1962; Higler & Duffels 1965). ® Z-Holland: Ouddorp,
Kwade Hoek, Brielse Meer (Krebs 1982). ® Zeeland:
Zierikzee (Fokker 1891; Duffels 1962), Yer-seke (Duffels
1962); Schouwen-Duiveland, 8 localities, Tholen, 1 loca-
lity, Noord-Beveland, 2 localities, Walcheren, 1 locality,
Zuid-Beveland, 5 localities, Zeeuws-Vlaanderen, 2 locali- Fig. 4. Sigara selecta.
ties (Krebs 1982).
Unpublished records. — © Friesland: Terschelling,
Striep, 22-24.viii.1962, J. H. Woudstra (W, BE). ® Zee-
land: Ellewoutsdijk, 11.vii.1987, B. Aukema.
24. Sigara hellensii (fig. 5)
References. — Snellen van Vollenhoven (1878a: 357);
Reclaire (1932: 230; 1936: 257; 1943: 121; 1948: 52);
Nieser (1982: 69).
Published records. — ® Gelderland: Leuvenum, Nun-
speet (Reclaire 1932); Arnhem, Warnsborn (Reclaire
1943); Hierdense Beek, Ellecom, Middachten (Nieser
1969; 1982). ® N-Brabant: Breda (Snellen van Vollenho-
ven 1878). ® Limburg: Belfeld (Reclaire 1936); Maasniel
(Reclaire 1948).
Unpublished records. - ® Gelderland: Leuvenum,
Hierdense Beek, 25-26.vii. 1961 & 3.viii. 1961, J. Beernink
& J. H. Woudstra; Oosterbeek, Hemelse Berg, 6.iv.1960,
R. H. Cobben.
32. Sigara iactans
References. - Cuppen (1988).
Published records. - ® Groningen: Midwolda (Cuppen
1988).
Unpublished records. — ® Groningen: Midwolda, 8 &
10.vi.1988 (7 À), 18.x.1988 (16 4), B. Aukema; Winscho-
ten, 8.vi.1988, 1 8, B. Aukema.
33. Sigara longipalis (fig. 6)
References. — Nieser (1982: 71).
Published records. — ® Overijssel: Oud-Ootmarsum
(Nieser). ® N-Brabant: Drunen, 15.vii.1981; Kaatsheu- Fig. 5. Sigara hellensti.
AUKEMA: Checklist Hemiptera-Heteroptera 37
vel, 18.viii.1981; De Moer, 12.viii.1981 (Nieser 1982;
IETS);
Unpublished records. - ® Groningen: Beerta,
8.vi.1988, 1 6, B. Aukema. © Drenthe: Beilen, Hijken,
14.iv.1974, 1 6, E. J. van Nieukerken (RML). ® Overijs-
sel: Ootmarsum, 15.vii.1974, 1 & at light, B. Aukema. ®
Utrecht: Eemnes, 14.ix.1988, R. Beenen (N). ® N-Bra-
bant: Hilvarenbeek, Beeksche Bergen, 20.vii.1985, 6 8, J.
Sparreboom (RML).
37. Naucoris maculatus (fig. 7)
References. - Reclaire (1932: 225; 1940: 114); Grave-
stein (1942: lviii); Nieukerken (1972).
Published records. - ® Utrecht: Vinkeveen (Grave-
stein 1942). e Z-Holland: Nieuwkoop (Van Nieukerken
1972). © N-Brabant: Oisterwijk (Reclaire 1932); Berli-
cum (Reclaire 1940).
Unpublished records. - ® Z-Holland: Noorden, De
Haeck, 26.iv.1969, N. Nieser (!EIS); Ibid, 10 &
25.viii.1974, B. Aukema; Ibid., 1.ix.1974, B. Aukema & J.
Beernink; Tienhoven, 13.x.1984, F van Nunen & R.
Beenen (P).
38. Aphelocheirus aestivalis (fig. 8)
References. - Reclaire (1932: 225; 1948: 51); Moller
Pillot (1971: 140).
Published records. — ® Overijssel: Oldenzaal (Reclaire
: ; re 1932). @ N-Brabant: Gilze-Rijen, Aa-gebied (Reclaire
Fig. 6. Sigara longipalis. 1932); Boxtel (Reclaire 1948); Dommelen (Moller Pillot
1971). ® Limburg: Mook, Heijen, Afferden (Reclaire
1932).
Unpublished records. — © N-Brabant: Spoordonk,
IF Beerze, 26.viii.1964, N. Nieser (!EIS).
Pe?
A 44. Notonecta reuteri (fig. 9)
CE
References. — Schober & Wassenaar (1980); Nieser
A (1982: 56).
Ô Note. - The specimens from Hoog-Buurlo, Gerritsfles
(20-23.vii. 1918, 2 9, M. Weber, coll. ZMA) recorded in
Reclaire (1932: 227) as N. lutea belong to this species.
Published records. - © Friesland: Terschelling, Vlie-
land (Schober & Wassenaar 1980). ® Drenthe: Beilen,
Dwingeloo, Havelte (Schober & Wassenaar 1980). ® Gel-
: derland: Hoog-Buurlo, Gerritsfles (Schober & Wassenaar
@-®@ | 1980); Buren (Nieser 1982).
ee | Unpublished records. — © Friesland: Terschelling,
Kroonpolders, 5.viii.1970, G. Visser (W). ® Overijssel:
Denekamp, Bergvennen, 2.vii.1970, 2 6, B. Aukema. ®
Drenthe: Ruinen, Kraloërplas, 22.ix.1984, 3 8, 4 9,
5 | 23.ix.1984, 1 &, 1 9, B. Aukema; Dwingeloo, IJsbaan,
| 23.ix.1984, 11 6, 2 9, 12.x.1984, 4 3, B. Aukema; Wijster,
| Meeuwenven, 2.x.1984, 2 g, 3 9, B. Aukema.
52. Velia saulii
References. - Van Nieukerken (1976).
Published records. - ® Limburg: Bunde, Cotessen,
Epen, Wittem (Van Nieukerken 1976).
Unpublished records. - ® Gelderland: Winterswijk,
Bekendelle, 5.v.1980, 1 9, K. den Bieman. ® Limburg:
Epen, Geul & Beversbeek, 4 & 5.v.1980, in large numbers,
Fig. 7. Naucoris maculatus. B. Aukema.
38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Fig. 8. Aphelocheirus aestivalis. Fig. 9. Notonecta reutert.
Fig. 10. Microvelia buenoi. Fig. 11. Gerris lateralis.
AUKEMA: Checklist Hemiptera-Heteroptera 39
53. Microvelia buenoi (fig. 10)
Synonym: M. #mbricola Wróblewski (Kanyukova 1986).
References. - Higler (1962, as M. #mbricola; 1967a);
Laeijendecker & Nieser (1971: 11, as zmbricola), Nieu-
kerken (1972, as umbricola); Nieukerken & van Tol
(1972, as umbricola); Van der Velde (1973, as #mbricola);
Cuppen (1978, as wmbricola), Giesen & Geurts (1985, as
umbricola).
Published records. - © Overijssel: Vollenhove, Dwars-
gracht (Higler 1962); Belt Schutsloot (Cuppen 1978). ®
Gelderland: Winterwijk (Laeijendecker & Nieser 1971);
Nijmegen, Hatertse- and Overasseltse vennen (Cuppen
1978); Doesburg, Eldrik, Gendringen, Hummelo, Hoog-
Keppel, Silvolde (Giesen & Geurts 1985). © N-Holland:
Kortenhoef (Higler 1962). ® Z-Holland: Nieuwkoop
(Nieukerken 1972); Schipluiden (Van der Velde 1973);
Schoonrewoerd, Everdingen (Cuppen (1978). ® Lim-
burg: Nederweert, Groote Peel (Higler 1967a; Nieuker-
ken & van Tol 1972); Meerlo-Wanssum, Geijsteren,
Horst, Mariapeel, Helden (Cuppen 1978).
Unpublished records. - ® Overijssel: Kalenberg, De
Weerribben (J. H. Woudstra; B. Aukema). ® Gelderland:
Driel (S. van Heijnsbergen, W); Wamel, Korenburger-
veen (K. den Bieman); Hemmen (B. Aukema); Buren (N.
Nieser et al., 'EIS). © Utrecht: Leersum (K. den Bieman);
Rhenen, Blauwe Kamer (B. Aukema); Westbroek (K. W.
R. Zwart, P). ®© N-Holland: Kortenhoef (P. van der Wiel,
ZMA; R. H. Cobben; J. H. Woudstra). ® N-Brabant:
Woudrichem, Loevestein (R. H. Cobben); Zundert (J. H.
Woudstra); Asten, Capelle, Drunen, Udenhout, Wern-
hout (N. Nieser et al., !EIS); Liessel (P. Poot). ® Limburg:
Leeuwen (P. van der Wiel, ZMA).
Fig. 12. Saldula scotica.
54. Microvelia pygmaea
References. - Reclaire (1940: 114)
Published records. - ® Limburg: Afferden, 14.ix.1935,
1 9, A. Reclaire (ZMA) (Reclaire 1940).
Unpublished records. - ® N-Brabant: Deurne, Mari-
apeel, Helenavaart, 30.ix.1983 (apterous @), 26.ix.1986
(apterous 9) & 26.ix.1987 (2 apterous 4), J. G. M. Cup-
pen; Ibid., Helenaveen, Helenavaart, 3.x.1986 (apterous
Q), J. G. M. Cuppen.
59. Gerris lateralis (fig. 11)
References. - Reclaire (1932: 217, as G. asper Fieber);
Cobben (1946: 58, as asper); Laeijendecker & Nieser
(1971: 11).
Published records. — © Gelderland: Winterswijk,
Meddo (Laeijendecker & Nieser 1971). ® N-Brabant:
Uden (Reclaire 1932). ® Limburg: Weert (Reclaire
1932); St. Odiliënberg (Cobben 1946).
Unpublished records. — ® Overijssel: Kalenberg,
Weerribben, 19.iv.1984, B. Aukema. ® Gelderland: Sta-
verden, 6.v.1969, micropterous 9, P. H. van Doesburg
(RML); Winterswijk, 15.iv.1979, micropterous 6, Seijs-
termans (ZMA). ® N-Brabant: Liempde, De Geelders,
9.x.1965, 13 & 15.v.1966, J. H. Woudstra; ibid., 17 &
28.vii.1966, B. Aukema; Best, 16.v.1985, 1 macropterous
9, B. Aukema.
Fig. 13. Saldula arenicola.
40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
67. Salda morio
References. — Reclaire (1932: 221).
Published records. — © Overijssel: Deventer,
20.vi.1910, macropterous 9, Klaassen (ZMA); Dene-
kamp, viii.1916, brachypterous 9, Koperberg (ZMA);
ibid., v.1918, Sth instar larva, D. MacGillavry (ZMA).
Present status. - Not recorded since 1918.
68. Salda muelleri
References. - Brakman (1960: 23).
Published records. — ® Zeeland: Hoofdplaat, 24.v-
28.vi.1959, P. J. Brakman, in large numbers, 2 macro-
pterous Q included (RML, ZMA, LUW, AU, P).
Unpublished records. — ® Friesland: Ameland, Oerd,
4.vii-18.viii.1985 & 31.viii.1986, Th. Heijerman; Ter-
schelling, Lies, 11.vii.1988, 1 9 & 12.vii.1988, 1 4, B.
Aukema.
70. Saldula scotica (fig. 12)
References. — Fokker (1884: 31; 1891: 358); Reclaire
(19825 2215 Osis Zl),
Present status. — Locally common in the southeast
(Limburg, Noord-Brabant and Gelderland) on stony ri-
versides.
71. Saldula arenicola (fig. 13)
References. — Fokker (1891: 358); Reclaire (1932:
223); Cobben (1950: 83; 1957: 255).
Published records. — © Friesland: Terschelling,
vii.1908 (1 8) & viii.1912 (2 @, 1 9), D. MacGillavry
(ZMA). ® Overijssel: Oldenzaal, vii.1906, 1 9, D. Mac-
Gillavry (ZMA). ® Gelderland: Velp, vii.1908, 1 4, D.
MacGillavry (ZMA). ® Z-Holland: Hoek van Holland, v,
E. Everts (RML).
Present status. - Locally common on sandy riversides
in the southeast (Limburg, Noord-Brabant, Gelderland
and Utrecht), elsewhere occasionally on sandy shores
along isolated smaller waterbodies.
72. Saldula c-album
References. - Cobben (1950: 83; 1957: 253).
Published records. - ® Limburg: Plasmolen,
22.vii.1949, 1 9, RH. Cobben (Cobben 1950); Eijsden,
vii, 1 6, E. Everts (RML) (Cobben 1957).
Unpublished records. - © Limburg: Epen, Cottessen,
in large numbers along De Geul, 8.viii.1983 &
16.viii.1986, Th. Heijerman; Ibid., 22.viii.1987, B. Au-
kema.
73. Saldula fucicola (fig. 14)
References. - Cobben (1950: 83, as S. vestita Douglas
& Scott; 1957: 252-253).
Present status. - Widespread and common along river-
sides and in the recently reclaimed Flevopolders.
74. Saldula melanoscela (fig. 15)
References. — Fokker (1884: 131, as Salda melanoscela;
1891: 358); Reclaire (1932: 222); Woudstra (1971).
Published records. — ® Friesland: Terschelling, Bos-
plaat (Woudstra 1971). ® Gelderland: Vorden (Fokker
Fig. 14. Saldula fucicola.
Fig. 15. Saldula melanoscela.
AUKEMA: Checklist Hemiptera-Heteroptera 41
1884). © Limburg: Eijsden (Fokker 1891), Valkenburg:
(Reclaire 1932).
Unpublished records. — © Friesland: Terschelling, Bos-
plaat, 19.viii.1971 & Ibid., Noordvaarder, 2.ix.1984, J. H.
Woudstra; Schiermonnikoog, Strandvlakte, 17.v, 11 &
12.vii and 15.ix.1972, A. Littel (AU, CB). ® Groningen:
Lauwersmeer, Ulrum, 11.iv & 10.v.1981, B. Aukema.
75. Saldula opacula (fig. 16)
References. — Reclaire (1940: 114); Cobben (1957:
253-255; 1959: 310-311).
Note. - The record from Zeeburg in Reclaire (1932)
concerns $. pallipes (Reclaire 1948: 51).
Published records. — ® Utrecht: Leersum (Reclaire
1940). ® Z-Holland: Oostvoorne (Cobben 1957). ® Lim-
burg: Heel, Horn (Cobben 1957).
Unpublished records. - ® Friesland: West-Vlieland, 8-
9.viii.1956, W. H. Gravestein (ZMA); Terschelling, Grilt-
jeplak, 24.viii.1954 (CB). ® Drenthe: Assen, 28.vi.1930,
1 9, H. C. Blöte (RML); Dwingeloo, 1.ix.1932, 1 9
(RML); Ibid., 2.vii.1967, S. van Heijnsbergen (W); Wij-
ster, IJsbaan, 6-12.ix.1975 & 15.iv.1981, B. Aukema; Ibid.,
Taaiveen, 26.iv.1976, B. Aukema; Ibid., Vossenberg,
24.vii.1984, B. Aukema; Spier, Kibbelhoek & Moddergat,
17.1v.1977, B. Aukema; Dwingeloo, Diepveen,
16.vi.1977, B. Aukema; Norg, Norgerholt, 24.vii.1977, B.
Aukema. ® Overijssel: Diepenveen, 13-14.vii.1967, 1 Q,
J. J. Meurer (ZMA). ® Gelderland: Delense Zand,
23.11.1952, 1 g, (CB); Gerritsfles, 16.iv.1959 (CB); Ha-
tertse Vennen, 25.x.1964, S. J. van der Molen (CB); Ton-
geren, 16.iv.1983, K. den Bieman (HE, BI). ® N-Holland:
Texel, de Geul, 29. viii.1974, 1 6, B. Aukema. © Limburg:
Horn, 6.iv.1963, R. H. Cobben.
Fig. 16. Saldula opacula.
78. Saldula palustris (fig. 17)
References. - Cobben (1957: 255).
Present status. - Widespread, coastal.
81. Micracanthia marginalis (fig. 18)
References. - Fokker (1899: 35, as Salda marginalis);
Cobben (1957: 255-256, as M. imitator (L.); 1959: 311-
312).
Note. - The record of Denekamp (Reclaire 1932: 223)
concerns S. saltatoria (Reclaire 1950: 22).
Published records. - ® Gelderland: Lochem, Winters-
wijk (Fokker 1899). ® Limburg: Horn (Cobben 1957).
Unpublished records. — © Friesland: Terschelling,
Hoorn, 12-13.viii.1965, J. H. Woudstra, Ibid,
23.viii.1965, J. Beernink; Ibid, Oosterend, 11-
19.viii. 1966, B. Aukema; Ibid., 13.vii. 1969, J. H. Woud-
stra. ® Gelderland: Staverden, 22.viii.1981, A. C. & W. N.
Ellis (ZMA). ® N-Brabant: Rul, 8.vi.1959, J.H. Woudstra
(W, BE, P).
84. Chartoscirta elegantula
References. - Reclaire (1932: 224; 1948: 51); Cobben
(1957: 256-257).
Published records. - © N-Holland: Diemen (Reclaire
1932). ® Limburg: Swalmen (Reclaire 1948); St. Odilién-
Fig. 17. Saldula palustris. berg (Cobben 1957).
42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Present status. - Not recorded since 1958: St. Odilién-
berg, 16.viii, R. H. Cobben (ZMA).
88. Acalypta marginata (fig. 19)
Note. — Older data on A. marginata, A. nigrina and A.
platycheila are not reliable since these species were often
misidentified in the past.
References. - Reclaire (1932: 127; 1936: 249; 1948:
43); Aukema (1983).
Published records. - © Limburg: Valkenburg (Reclaire
1936); Terblijt (Reclaire 1948); Stokhem (Aukema
1983).
Unpublished records. - ® Gelderland: Schaarsbergen,
21.vii.1955, R. H. Cobben. ® Utrecht: Leersum,
12.v.1979, 1 g, K. den Bieman. ® N-Holland: Hilversum,
18.v.1985, R. Ph. Jansen (ZMA, JA); Ibid, 2v &
10.v.1986, J. H. Woudstra, 5.v.1986, B. Aukema &
31.v.1986, K. den Bieman. ® Limburg: Maastricht,
vi.1894 (RML) (Reclaire 1932, as platychila); Linne,
25.v.1944, 1 g, R. H. Cobben (Reclaire 1948, as nigrina).
89. Acalypta musci
References. — Reclaire (1932: 126; 1934: 51).
Published records. — Valkenburg, Houthem (Reclaire
1932, 1934).
Present status. - Collected between 1923 and 1943 on
the same locality.
90. Acalypta nigrina (fig. 20)
Note. - older data are unreliable (see marginata and
platycheila).
References. - Reclaire (1932: 127; 1936: 249-250).
Published records. - ® Gelderland: Groesbeek, Leuve-
num (Reclaire 1936).
Unpublished records. - ® Drenthe: Kraloo,
22.vii.1959, 22.vi.1960, 28.vi & 2.viii.1961, 25.vii. 1962,
17.vi & 22.vii.1964, 11 & 18.viii.1965 & 24.viii.1966, P. J.
den Boer (BSW). © Gelderland: Eerbeek, v.1916, 1 9, D.
L. Uyttenboogaart (ZMA) (Reclaire 1932, as marginata);
Wageningen, Oostereng, ix.1949, 1 9, J. van der Drift
(RML). ® Utrecht: Driebergen (Reclaire 1932, as nigrina
and marginata). ® N-Holland : Hilversum, 13.vi.1965,
M. J. Gijswijt (W); Ibid., 22.viii-4.x.1986, R. Ph. Jansen
(JA, W, ZMA).
92. Acalypta platycheila (fig. 21)
Note. — older data are unreliable (see marginata and
nigrina).
References. - Reclaire (1932: 127; 1943: 110; 1948:
43).
Published records. — ® Drenthe: Assen (Reclaire
1948).
Unpublished records. — ® Friesland: Terschelling, Bos-
plaat, 15.vii.1958, W. H. Gravestein (ZMA); Ameland,
Oerd, 18.viii.1985, Th. Heijerman. ® Drenthe: Lhee-
broek, 29.vi, 13 & 20.vii.1960, P.J. den Boer (BSW, ZMA);
Wijster, IJsbaan, 11.vi.1976, B. Aukema. ® Gelderland:
Winterswijk, 22.ix.1941, A. Reclaire & Pvan der Wiel
(ZMA) (Reclaire 1943, as nigrina); Bennekom, 16-17.vi
& 8.vii.1956, R. H. Cobben.
Fig. 18. Micracanthia marginalis.
Fig. 19. Acalypta marginata.
AUKEMA: Checklist Hemiptera-Heteroptera 43
97. Galeatus maculatus
References. - Fokker (1884: 124).
Published records. - ® Gelderland: Wageningen, 1 3,
J. Ritzema Bos (RML) (Fokker 1884).
Present status. - Not recorded since 1884.
100. Lasiacantha capucina
References. - Reclaire (1936: 250).
Published records. - © Limburg: Bemelen, 26 &
27.iv.1935, M. van den Boorn & A. M. Scholte (Reclaire
1936).
Unpublished records. - © Limburg: Bemelen, 2.v.1935
& 16.iv.1936, M. van den Boorn (NMM); Ibid., 1.vi.1944,
M. van den Boorn (P); Ibid, 6.viti.1948, 30.vi &
23.vit.1950, P. van der Wiel (ZMA); Ibid., 30.vi.1950, C.
Berger (P).
Present status. - Not recorded since 1950.
101. Tingis reticulata
References. - Fokker (1884: 124, as Monanthia ciliata
Fieber); Reclaire (1932: 131-132; 1936: 250; 1948: 43);
Cobben (1958: 16)
Published records. - © Gelderland: Wageningen (Cob-
ben 1958). © N-Holland: Bloemendaal (Fokker 1884). ®
Z-Holland: Meijendel (Reclaire 1932); Wassenaar (Re-
claire 1936). ® N-Brabant: Breda, Liesbos (Reclaire
1948).
Fi AET Se Unpublished records. - ® N-Holland: Santpoort, Duin
ig. 20. Acalypta nigrina. en Kruidberg, 18.v, 27.vi, 3 & G.viii.1974, B. Aukema,
23.v.1974 & 19.v.1975, J. H. Woudstra & 21.11.1976 &
17.iv.1981, B. Aukema & J. H. Woudstra. ® Z-Holland:
Meijendel, Bierlap, 3.x.1944, 1 9, M. van den Boorn (P).
102. Tingis pilosa
References. — Reclaire (1932: 132; 1940: 109); Blöte
(1954: 84).
Published records. - ® Z-Holland: Noordwijk (Re-
claire 1932). ® Limburg: Schin op Geul (Reclaire 1940);
Eijsden (Blöte 1954).
Unpublished records. - ® Limburg: Cottessen, 26 &
30.viii. 1968, J. H. Woudstra; Ibid, 12 & 17.viii.1971,
10.1x.1973, B. Aukema, 17.viii.1974, B. Aukema & J. Beer-
nink; Stramproij, 11.vi.1984, K. den Bieman.
105. Tingis crispata
References. - Aukema (1976).
Published records. - © Limburg: Maastricht, St. Pie-
tersberg (Aukema 1976).
Unpublished records. - ® Limburg: Maastricht, St.
Pietersberg, 29.iv, 8-11.viii & 2.ix.1983, B. Aukema; Ibid.,
15.x.1984, B. van Aartsen (ZMA). ® N-Brabant: Best,
4.vi.1983 & 19.v.1984, B. Aukema & J. H.Woudstra,
16.v.1985, B. Aukema & H. W. van der Wolf.
107. Physatocheila costata
References. — Cobben (1958a: 10-12, as gwadrimacu-
lata).
Published records. - ® Limburg: Haelen (Nunhem),
Maasniel, Susteren (Cobben 1958).
Present status. - Not recorded since 1951: Susteren,
Fig. 21. Acalypta platycheila. 2.vill, R. H. Cobben (CB, AU).
44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Fig. 22. Physatocheila dumetorum. Fig. 23. Physatocheila smreczynskit.
Fig. 24. Deraeocoris punctulatus. Fig. 25. Deraeocoris annulipes.
AUKEMA: Checklist Hemiptera-Heteroptera 45
et 108. Physatocheila dumetorum (fig. 22)
Present status. - Formerly a widespread species, but
recorded only twice since 1957 from the province of
Zeeland: Oostkapelle, 8.ix.1962 & Vrouwenpolder,
14.ix.1962, P. J. Brakman (P).
109. Physatocheila smreczynskii (fig. 23)
References. - Cobben (1958: 10-12).
Note. - Most old records of P. quadrimaculata (Reclaire
1932, 1934, 1936, 1943, 1948 & 1951) concern this spe-
cies.
Present status. - Widespread.
110. Oncochila simplex
References. - Reclaire (1932: 134); Cobben (1958a: 12-
14).
Published records. - ® Overijssel: Deventer, viii.1912,
Klaassen (Reclaire 1932). ® Limburg: Rijkel, 7.vi.1953,
1 9, W. H. Gravestein (ZMA); Ibid., 2-7.viii.1955, in
numbers, R. H. Cobben (Cobben 1958a).
Unpublished records. — © Gelderland: Winssen,
31.v.1981, 1 &, Th. Heijerman; Ibid., 14.v.1982, 1 6, K.
den Bieman; Ibid, 29.v & 13.vii.1985, 19.v.1986,
el 24.v.1987 & 2.v.1988, B. Aukema.
119. Myrmedobia distinguenda
Fig. 26. Deraeocoris scutellaris. References. - Fokker (1899: 35); Reclaire (1932: 156).
Published records. - © Gelderland: Rheden, De Steeg
(Fokker 1899); Putten, Leuvenum (Reclaire 1932).
Present status. - Not recorded since 1913: Leuvenum,
9.vi, 1 9, D. MacGillavry (ZMA).
123. Bothynotus pilosus
References. - Fokker (1891: 359); Reclaire (1936: 254;
1948: 47).
Published records. - ® Utrecht: Den Dolder (Reclaire
1936). ® Gelderland: Groesbeek (Reclaire 1936). ® N-
Brabant: Breda (Reclaire 1948). ® Limburg: locality un-
known (Fokker 1891).
Unpublished records. - © N-Brabant: Liempde, De
Geelders, 14.vi.1966, 1 macropterous @, at light, B. Au-
kema.
125. Deraeocoris punctulatus (fig. 24)
References. - Reclaire (1932: 176; 1936: 254; 1943:
115; 1948: 47; 1951: 15); Meurer (1957: 95); Brakman
(1960: 21).
Present status. — Formerly widespread, but recorded
only once since 1960: Limburg, Vlodrop, 20.viii. 1968, 1
6, P. J. Brakman (P).
126. Deraeocoris annulipes (fig. 25)
References. - Cobben (1958a: 21-22).
Present status. - Widespread, but rare.
128. Deraeocoris flavilinea f. nov. spec.
Since 1985 recorded from Arnhem, Nijmegen and Wa-
geningen from common maple (Acer campestre L.) and
Fig. 27. Oncotylus punctipes. sycamore (A.pseudoplatanus L.).
46 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Records. — ® Gelderland: Wageningen, 13-20.vi.1985,
16.vi-6.vii.1987 & 2.vi.1988, B. Aukema (6.vii.1987: B.
Aukema & J. H. Woudstra); Ibid., 5.vi.1986, K. den Bie-
man; Arnhem, 16.vi.1985, B. Aukema; Nijmegen, 16.vi-
5.vii.1985, 22.vi.1986, 28.vi-11.ix.1987 & 21.vi-1.vii.1988,
B. Aukema.
131. Deraeocoris scutellaris (fig. 26)
References. — Fokker (1899: 35, as Capsus scutellaris).
Published records. — ® Limburg: Venlo (Fokker 1899).
Unpublished records. - ® Drenthe: Hooghalen, vi-
vii 1944, E. A. M. Speijer (RML); Kraloo, 17.vii.1966, 1
®, P.J. den Boer (BSW); Westerbork, 3.x.1968, 1 9, J. B.
Wolschrijn (ZMA); Wijster, 2, 12-13 & 28.vii.1975, 15-
16, 20.vi & 26.vii.1978, B. Aukema; Nuil, 6.viii.1975, B.
Aukema; Beilen, 27.vi.1983, 1 9, B. van Aartsen (ZMA);
Dwingeloo, 25.viii.1984, B. van Aartsen (W). © Overijs-
sel: Diepenveen, 13-14.vii.1967, J. J. Meurer (ZMA). ©
Gelderland: Terlet, 1.1x.1979, B. Aukema: Ibid,
20.vi.1982, B. van Aartsen (ZMA). Ede, Ginkelse Heide,
3.vii.1985, 1 g, B. Aukema.
136. Oncotylus punctipes (fig. 27)
References. — Fokker (1891: 360); Reclaire (1932:
200); Gravestein (1949: xxiv).
Published records. - € Overijssel: Denekamp (Grave-
stein 1949). ® Gelderland: Arnhem (Fokker 1891);
Warnsveld (Reclaire 1932).
Unpublished records. - ® Overijssel: Vasse, 2.vii.1970,
B. Aukema; Ibid, 8.vii. 1972, B. Aukema & J. H. Woudstra;
Ibid, 14.vii.1972 & Ootmarsum, 5.vii.1970, J. H. Woud-
Stra. © Limburg: Swalmen, 1.viii.1980, B. Aukema;
Venlo, 3.viii. 1983, J. H. Woudstra & B. Aukema.
157. Oncotylus viridiflavus
References. — Brakman (1951); Cobben (1953: 194-
196; 1958: 20).
Published records. - © Zeeland: Nieuw & St. Joosland
(Brakman 1951). ® Gelderland: Valburg (Cobben 1953);
Wageningen (Cobben 1958).
Unpublished records. - ® Zeeland: Vrouwenpolder,
15.viii.1955, P. J. Brakman (P); Nisse, 22.viii.1960, 1 é,
P. J. Brakman (RML).
159. Conostethus roseus (fig. 28)
References. — Snellen van Vollenhoven (1878a: 162-
163, as Lopus subpatellatus nov. spec); Fokker (1885:
70); Aukema (1988).
Published records. - ® Drenthe: Noordsleen (Aukema
1988). ® Flevoland: Biddinghuizen (Aukema 1988). e
Gelderland: Velp (Snellen van Vollenhoven 1878); Ede
(Fokker 1885; Aukema 1988); 't Harde, Hulshorst, Koot-
wijk, Otterlo, Hoge Veluwe, Tongeren (Aukema 1988). ®
N-Holland: Hilversum (Aukema 1988). ® Limburg: Vel-
den (Aukema 1988).
Unpublished records. — © Overijssel: Ommen,
3.vi.1987, B. Aukema & D. J. Hermes; Ibid., 4.vi.1988, B.
Aukema & G. Morseld. ® Gelderland: Wekeromse Zand,
1.v.1988, larvae, B. Aukema.
Fig. 28. Conostethus roseus.
Fig. 29. Conostethus venustus.
AUKEMA: Checklist Hemiptera-Heteroptera 47
140. Conostethus venustus (fig. 29)
References. - Aukema (1988). '
Published records. - ® Utrecht: Remmerden, Rhenen
(Aukema 1988). ® Gelderland: Heelsum, Loenen aan de
Waal (Aukema 1988). ® Zeeland: Veere, Vlissingen,
Westkapelle, Wissenkerke (Aukema 1988). ® Limburg:
Broekhuizenvorst, Lomm, Lottum, Maastricht, Meche-
len, Oost-Maarland, Schandelo, Strucht, Tienray, Velden,
Venlo, Wijlre (Aukema 1988).
Unpublished records. - ® Gelderland: Berg en Dal,
22.vi & 21.viii.1986, K. den Bieman; Ubbergen, Ooijsche
Graaf, 16.vii.1987, B. Aukema; Millingen aan de Rijn,
16.vii.1987, B. Aukema; Doorwerth, Duno, 26.vi &
9.vii.1988, B. Aukema; Wageningen, 15.viii.1988, B. Au-
kema.
141. Hoplomachus thunbergi (fig. 30)
References. - Snellen van Vollenhoven (1878a: 216-
217, as Lygus thunbergti), Fokker (1885: 70); Reclaire
(1932: 200-201; 1940: 112; 1943: 117).
Present status. — Formerly widespread, but not recor-
ded since 1951: N-Brabant, Wouw, 17.vi.1951, R. H.
Cobben (LUW).
145. Amblytylus brevicollis
References. - All records in Fokker (1885: 69) and
Reclaire (1932: 202) concern A. nasutus (Reclaire 1936:
Fig. 30. Hoplomachus thunbergi. 255-256).
Unpublished records. — © Gelderland: Wolfheze,
8.vii.1882, 1 9, A. J. F Fokker (RML); Vierhouten,
23.vii.1962, 1 9, J. Beernink (det. E. Wagner).
148. Macrotylus solitarius (fig. 31)
References. — Reclaire (1943: 118); 1948: 49; 1951:
19); Blöte (1954: 83); Cobben (1958: 20).
Published records. - © Limburg: Bemelen (Reclaire
1943); Maasniel, Boukoul (Reclaire 1948); Geulhem (Re-
claire 1951); Maastricht (Blöte 1954); Asselt (Cobben
1958).
Unpublished records. - ® N-Brabant: Liempde, De
Geelders, 23.vii.1966, B. Aukema; Best, De Mortelen,
31.vii.1977, H. W. van der Wolf (ZMA). ® Limburg:
Berg, 16.viii.1961, J. H. Woudstra; Bemelen, 20.vii.1965,
J. H. Woudstra; Vijlen, 21.vii.1965, J. H. Woudstra;
Wijlre, 6 & 12.viii. 1983, B. Aukema.
151. Tytthus geminus f. nov. spec.
Records. - ® Drenthe: Wijster, IJsbaan, 12.1x.1975, 1
brachypterous 9, B. Aukema.
152. Tytthus pygmaeus (fig. 32)
References. - Fokker (1885 : 67, as Cyrtorrhinus pyg-
maeus).
Published records. - © Zeeland: Renesse (Fokker
1885).
Unpublished records. — ® Friesland: Vlieland.
viii. 1956, W. H. Gravestein (ZMA); Terschelling, Ooster-
end, 27.vii.1958 & 5-12.vii.1959, W. H. Gravestein
(ZMA); Ibid., 19.viii.1963 & 12.viii.1964, J. H. Woudstra;
Fig. 31. Macrotylus solitarius. Ibid., 12.viti.1964, J. Beernink. Ibid, Noordvaarder,
48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Fig. 32. Tytthus pygmaeus. Fig. 33. Psallus pseudoplatani.
Fig. 34. Psallus wagneri (males). Fig. 35. Psallus confusus (males).
AUKEMA: Checklist Hemiptera-Heteroptera 49
23.viii.1963, J. H. Woudstra; Ibid., Hoorn, 15.viii.1969;
Ibid., Koegelwieck, 2-3.viii.1987, B. Aukema; Ibid., 12 &
14.vii.1988, B. Aukema & J. H. Woudstra. ® Drenthe:
Dwingeloo, 23.vi.1976, 1 9, B. Aukema; Norg, Norger-
holt, 24.viii.1977, B. Aukema. ® Overijssel: Kalenberg,
Weerribben, 2.viii.1965, 1 9, J. Beernink. ® Flevoland:
Lelystad, 1.vii.1986, 1 9, B. Aukema. ® Gelderland: Hoge
Veluwe, 3.viii.1983, 1 6, R. Ph. Jansen. ® Zeeland: Vlis-
singen, 16.vii.1986 & Westkapelle, 25.vii. 1986, B. Au-
kema.
153. Brachyarthrum limitatum f. nov. spec.
Recorded since 1980 from aspen (Populus tremula L.).
Records. - ® Gelderland: Doorwerth, Duno, 26-27.vi
& 9.vii.1988, B. Aukema & G. Morseld. © Limburg:
Venlo, 17.vi.1980 & 10.vi.1982, B. van Aartsen (ZMA);
Ibid., 23-25.vi.1981, 13, 15 & 17.vi.1982, B. Aukema;
Lomm, Ravenvennen, 8-9.vi.1982, B. Aukema & J. H.
Woudstra; Lottum, 14 & 16.vi.1982, B. Aukema.
161. Psallus assimilis
References. - Aukema (1986).
Published records. - ® Limburg: Maastricht, Meche-
len, Wittem (Aukema 1986).
Present status. - Collected in 1985 only.
163. Psallus pseudoplatani (fig. 33)
Fig. 36. Psallus salicis. References. - Aukema (1986).
Published records. - ® Drenthe: Meppel (Aukema
1986). ® Gelderland: Arnhem, Ede, Nijmegen, Ooster-
beek, Wageningen (Aukema 1986). ® N-Holland : Zaan-
dam (Aukema 1986). ® Limburg: Eygelshoven, Maas-
tricht, Mechelen (Aukema 1986).
Present status. - Widespread, collected since 1976.
165. Psallus wagneri (fig. 34)
References. - Aukema (1981).
Present status. - Widespread.
167. Psallus confusus (fig. 35)
Synonym: P. diminutus auct. nec Kirschbaum (Rieger
1981).
References. - Aukema (1989).
Present status. - Widespread.
172. Psallus mollis
Synonym: P. diminutus Kirschbaum (Rieger 1981).
References. - Aukema (1989).
Published records. - © Overijssel: Delden (Aukema
1988). © N-Brabant: Breda (Aukema 1988).
Unpublished records. - ® Gelderland: Doorwerth, 26-
27.vi & 9.vii.1988, B. Aukema.
173. Psallus salicis (fig. 36)
Synonym: P. alnicola Douglas & Scott (Rieger 1981).
References. - Reclaire (1943: 118, as a/nicola); Cobben
(1958: 20, as alnicola).
Note. - Nearly all the older records of P. salicis (as P.
alnicola) are based on misidentifications.
Fig. 37. Atractotomus parvulus. Published records. - € Gelderland: Hall, 18.viii.1939,
50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Fig. 38. Plagiognathus litoralis. Fig. 39. Plagiognathus alpinus.
Fig. 40. Plagiognathus fulvipennis. Fig. 41. Chlamydatus evanescens.
AUKEMA: Checklist Hemiptera-Heteroptera Si
1 ®, A. Reclaire (ZMA) (Reclaire 1943). ® Limburg:
Melick-Herkenbosch, viii.1947 & viii.1952, R. H. Cobben
(Cobben 1958).
Unpublished records. - ® Friesland: Terschelling,
West-Terschelling, 30.viii.1986, J. H. Woudstra. ® Gel-
derland: Doorwerth, 22.viii.1985, B. Aukema; Renkum,
29.viii.1987, 1 9, K. den Bieman; Wageningen,
30.vii.1986, B. Aukema & J. H. Woudstra. ® Utrecht:
Loosdrecht, 22.vii.1959, J. H. Woudstra; Kamerik,
10.viii.1974, B. Aukema. ® N-Holland: Amsterdam, 1-
19.viti.1974, B. Aukema; Zaandam, 9.viii.1974 & 5-
6.viii.1975, J. Beernink; Westzaan, 10.viii.1986, J. H.
Woudstra. © Z-Holland: Noorden, De Haeck,
24.vii.1974, B. Aukema; Middelharnis, 27.viii.1968, 1 8,
P. Vroegindeweij (RML). © N-Brabant: Made,
29.viii.1987, 1 9, K. den Biemans; Liessel, 9.ix.1987, B.
Aukema.
180. Atractotomus parvulus (fig. 37)
References. - Reclaire (1943); Meurer (1957: 92).
Note. - The specimen recorded by Cobben (1958: 20)
belongs to A. magnicornis.
Published records. - ® Gelderland: Elden (Reclaire
1943). © Utrecht: Baarn (Reclaire 1943). © N-Holland:
Laren (Reclaire 1943); Heemstede (Meurer 1957). ©
Limburg: Mook (Reclaire 1943).
Unpublished records. - © Drenthe: Dwingeloo,
à i 19.vii.1949, W. H. Gravestein (ZMA). ® Gelderland:
Fig. 42. Monosynamma sabulicola. Schaarsbergen, viii.1968, H. Klomp et al. (CB). ® N-
Holland: Zaandam, 20.vii.1973 & 17.viii.1981, J. H.
Woudstra; Santpoort, 6.viii.1974, B. Aukema. © N-Bra-
bant: Herpen, 31.viii.1962, S. van Heijnsbergen (W).
182. Plagiognathus litoralis (fig. 38)
References. - Reclaire (1932: 210-211, as P. fusciloris);
Gravestein (195 la); Brakman (1960: 21).
Present status. - Widespread, coastal.
183. Plagiognathus alpinus (fig. 39)
References. - Fokker (1885: 71); Reclaire (1936: 256;
19571220)
Published records. - ® Limburg: Valkenburg (Fokker
1885); Geulle (Reclaire 1936); Geulhem, Wijnandsrade,
Haelen, St. Odiliënberg, Beek, Meerssen (Reclaire 1951).
Unpublished records. - ® Limburg: Camerig, Epen,
Vijlen (J. H. Woudstra); Slenaken (P. Poot); Cottessen (J.
H. Woudstra & B. Aukema); Holset, St. Geertruid (B.
Aukema).
186. Plagiognathus fulvipennis (fig. 40)
References. - Reclaire (1932: 211; 1940: 113; 1943: 19;
1951: 20).
Present status. - Many older records, but only one
recent capture: Ootmarsum, Springendal, 24.vii.1971, 2
6, J. H. Woudstra.
191. Chlamydatus evanescens (fig. 41)
References. - Cobben (1960: 195-205).
Fig. 43. Campylomma annulicorne. Present status. - Widespread.
52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
192. Monosynamma bohemanni
Synonym: M. nigritulum (Zetterstedt) (Josifov 1961).
194. Monosynamma sabulicola (fig. 42)
References. — Stichel (1956: 357, no records given).
Unpublished records. — ® Flevoland: Lelystad (H. Val-
lenduuk (ZMA); B. Aukema). ® Gelderland: Elden (A.
Reclaire A. J. Winkelman (ZMA)); Doornenburg, Epe,
Tolkamer, Valburg (B. Aukema); Wageningen (B. Au-
kema & J. H. Woudstra); Lienden (K. den Bieman). ®
Utrecht: Zeist (K. den Bieman). ® N-Holland: Hilver-
sum (Reclaire ZMA). ® Z-Holland: Gorinchem (P.
Poot). ® N-Brabant: Biesbos (B. v. Aartsen, M. Koning
(ZMA); R. H.Cobben). ® Limburg: Arcen (W. H. Gra-
vestein); Asselt (W. H. Gravestein; B. Aukema); Echt (P.
J. Brakman, P); Urmond (P. Poot).
195. Campylomma annulicorne (fig. 43)
References. — Reclaire (1946: 50; 1951: 20); Meurer
(1957); Cobben (1958: 21).
Published records. - € Utrecht: Vinkeveen (Reclaire
1946). © N-Holland: Edam (Reclaire 1946); Heemstede
(Meurer 1957). ® Limburg: Roermond (Reclaire 1950);
Boukoul (Cobben 1958).
Present status. - Widespread.
197. Criocoris crassicornis (fig. 44)
References. — Fokker (1886: 299); Reclaire (1932:
210); Cobben (1953: 196).
Note. - The specimen from Ommen (vi.1916, D. Mac-
Gillavry) mentioned by Reclaire (1932) as questionable,
could not be found in any collection.
Published records. - © Limburg: Nuth (Fokker 1886);
Haelen, Leeuwen, St. Odiliënberg (Cobben 1953).
Unpublished records. — © Gelderland: Bennekom,
9.viii.1986, K. den Bieman. ® Limburg: Epen, Onderste
Bos, 29.vii, H. C. Blöte (RML); Ibid., Bovenste Bos,
16.viii.1971, B. Aukema; Ibid., 17.viii.1974, B. Aukema &
J. Beernink.
198. Criocoris sulcicornis f. nov. spec.
Records. — © Zeeland: Valkenisse, 26.vii.1967, 1 9, B.
van Aartsen (ZMA).
201. Asciodema obsoletum (fig. 45)
References. — Reclaire (1934: 60; 1940: 130); Cobben
(1958: 21).
Published records. - ® Limburg: Spaubeek (Reclaire
1934); Epen (Reclaire 1940); Horn, Melick-Herkenbosch
(Cobben 1958).
Unpublished records. - ® Utrecht: Veenendaal,
30.vi.1988, B. Aukema. © Gelderland: Oosterbeek,
23.vi.1960, R. H. Cobben; Nijmegen, 28.vi & 5.vii.1985,
B. Aukema; Berg en Dal, 22.vi.1986, K. den Bieman. ®
Zeeland: Oostkapelle, 18.vii.1986, B. Aukema. © N-Bra-
bant: Waalre, Treeswijk, 2.viii. 1941, H. C. Blöte (RML).
© Limburg: Vilt, 22.vu.1965, J. H. Woudstra; Cottessen,
18.vii.1974, B. Aukema & J. Beernink; Herkenbosch,
Meijnweg, 31.vii.1980, B. Aukema.
Fig. 44. Criocoris crassicornis.
Fig. 45. Asciodema obsoletum.
AUKEMA: Checklist Hemiptera-Heteroptera 53
202. Hallodapus rufescens (fig. 46)
References. — Gravestein (1952a: cxviii); Cobben
(1958: 20); Woudstra (1959: 106).
Published records. - € Gelderland: Otterloo (Grave-
stein 1952a). © N-Holland: Heemskerk, Bakkum (Woud-
stra 1959). e Z-Holland: Katwijk (Cobben 1958).
Unpublished records. - € Drenthe: Wijster, vii. 1975,
1 macropterous 9, at light, B. Aukema, ® Overijssel:
Denekamp, Bergvennen, 29.vi.1970, B. Aukema. ® Gel-
derland: 't Harde, 5.viii.1962, J. H. Woudstra; Imbosch,
W.K. R. E. van Wingerden et al. (RINA). ® N-Holland:
Bakkum, 19.viii.1960, J. H. Woudstra & J. Beernink; Ibid.,
9.vii & 20.viii.1969, 3.vii.1965, J. H. Woudstra.
204. Cremnocephalus albolineatus
References. - Reclaire (1936: 255).
Published records. - ® Overijssel: Denekamp (Re-
claire 1936).
Unpublished records. - ® Drenthe: Spier, Kibbelhoek,
l.viii.1981 & 28.vii.1984, B. Aukema. € Overijssel: Weer-
selo, 20.vi.1952, W. H. Gravestein (ZMA); Lutterzand,
18.vi 1960, W. H. Gravestein (ZMA); Ootmarsum, Sprin-
gendal, 26-27.vi & 4.vii.1970, B. Aukema; Ibid., 5.vii.1970,
J. H. Woudstra; Stegeren, 27.vii.1986, H. Vallenduuk;
Ommen, Eerderveld, 4.vi.1988, larvae, B. Aukema.
206. Dicyphus constrictus
. References. - Reclaire (1932: 185).
Fig. 46. Hallodapus rufescens. Published records. - ® N-Holland: Wijk aan Zee,
viii.1910, 1 9, D. MacGillavry (ZMA) (Reclaire 1932).
Present status. - Not recorded since 1910.
214. Hypseloecus visci
References. — Arnoud (1958); Cobben & Arnoud
(1969).
Published records. - © Limburg: Voerendaal,
22.viii.1958, Br. Arnoud (Arnoud 1958); Mechelen,
15.vi.1960, larvae, Br. Arnoud (Cobben & Arnoud 1969).
Unpublished records. - ® Limburg: Geulle,
16.viii.1966, P. J. Brakman (RML) & P. Poot.
220. Halticus luteicollis
References. - Fokker (1891: 359); Reclaire (1936: 255;
1940: 112); Blöte (1950: 19).
Published records. - ® Limburg: Valkenburg (Fokker
1891); Bemelen (Reclaire 1936), Gronsveld, Schin op
Geul (Reclaire 1940); Maastricht (Blöte 1950).
Unpublished records. — © Limburg: Bemelen,
22.vi.1948, P. van der Wiel (P); Schin op Geul,
14.viii.1961 & 29.vii.1964, J. H. Woudstra; Vlodrop,
23.vii.1985, 1 9, B. van Aartsen (ZMA).
221. Halticus saltator
References. - Fokker (1891: 359, as H. erythrocepha-
lus); Reclaire (1932: 198; 1943: 117).
Published records. - ® Gelderland: Huissen (Reclaire
1932). ® N-Brabant: Cuyk (Fokker 1891). ® Limburg:
Well (Reclaire 1932). ® Zeeland: Hulst (Reclaire 1943).
Unpublished records. - © Limburg: Bemelen,
22.vi.1948, 1 9, P. van der Wiel (RML).
Fig. 47. Strongylocoris leucocephalus. Present status. - Not recorded since 1948.
54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
222. Strongylocoris leucocephalus (fig. 47)
References. — Snellen van Vollenhoven (1878a: 171-
172, as Halticus leucocephalus), Fokker (1885: 63); Re-
claire (1932: 197; 1936: 255; 1948: 49; 1951: 18); Cobben
(1958: 19).
Published records. - ® Gelderland: Wolfheze (Snellen
van Vollenhoven 1878); Ede, Oosterbeek, Vorden, Wage-
ningen (Fokker 1885); Bennekom (Cobben 1958). ®
Utrecht: Eemnes (Reclaire 1948); Rhenen (Cobben
1958). e N-Brabant: Breda (Snellen van Vollenhoven
1878). © Limburg: Epen (Reclaire 1936); Neer (Reclaire
1950).
Unpublished records. - ® Drenthe: Rolde, Balloérveld
(W. H. Gravestein, ZMA). ® Overijssel: Deventer
(ZMA), Eerde (RML). ® Gelderland: Garderen, Hoge
Veluwe, Warnsveld (ZMA); Groesbeek (RML, ZMA);
Wageningen (ZMA, CB); Doetinchem (FNM). ®
Utrecht: Amersfoort (RML). ® N-Brabant: Breda (RML,
ZMA). ® Limburg: Mook (ZMA); Rijckholt (NMM).
Present status. — Formerly widespread, but recorded
only once since 1956: Rolde, Balloérveld, 10.vi.1961, 1 9
W. H. Gravestein (ZMA).
226. Orthocephalus saltator
Synonym: Tamanini (1977) synonymized O. ferrari Reu-
ter with O. saltator.
227. Reuteria marqueti f. nov. spec.
Recorded since 1987 from elm (Ulmus sp.) and lime
(Tilia sp.).
Records. - © N-Holland: Naarden, 31.viii & 2.ix.1987,
31.viii.1988, J. H. Woudstra. ® Gelderland: Wageningen,
1, 4 & 27.ix.1987, 4.ix.1988, B. Aukema; Nijmegen,
11.ix.1987, B. Aukema.
229. Fieberocapsus flaveolus (fig. 48)
References. — Gravestein (1952a: cxviii); Brakman
(1960: 22).
Published records. - ® N-Holland: Texel, Slufter (Gra-
vestein 1952a). ® Zeeland: Hoek, Hoofdplaat (Brakman
1960).
Unpublished records. - ® Friesland: Ameland, Oerd,
31.vii.1967, B. Aukema; Terschelling, 10.viii.1953, P. E.
van Heerdt; Terschelling, Oosterend, 19.viii.1962, J. H.
Woudstra; Ibid., Bosplaat, 14-19.viii.1964, J. H. Woudstra
& J. Beernink; Ibid, 12 & 19.viii.1966, 6.vii.1967 &
9.viii.1970, J. H. Woudstra; Ibid., 2.viii.1987, B. Aukema;
Ibid, Noordvaarder, 14.viii.1965 & 25.viii.1984, J. H.
Woudstra; Ibid, Koegelwieck, 3.viii.1987, B. Aukema;
West-Vlieland, 12.viii.1956, W. H. Gravestein (ZMA,
W). © Z-Holland: Voorne, Rockanje, 1.vii.1970 (ZMA).
® Zeeland: Westkapelle, 23 & 25.vii.1986, B. Aukema.
234. Globiceps sphegiformis
References. - Fokker (1885: 67; 1891: 360); Reclaire
(1932: 190).
Published records. — ® Limburg: Valkenburg, vii, 1 9,
A.J. F. Fokker (RML) (Fokker 1885); Ibid., vii.1887, 1 Q,
E. Everts (RML) (Fokker 1891); Epen, 20.vii.1916, 1 &,
D. MacGillavry (ZMA) (Reclaire 1932).
Fig. 48. Fieberocapsus flaveolus.
Fig. 49. Heterocordylus leptocerus.
AUKEMA: Checklist Hemiptera-Heteroptera 55
Unpublished records. - ® Limburg: Epen, 5.vii.1924,
1 9, H. C. Blöte (RML).
Present status. - Not recorded since 1924.
235. Heterocordylus genistae f. nov. spec.
Note. - The records in Reclaire (1932: 195, Hilversum;
1934: 58, Doetinchem) concern Orthocephalus saltator
and H. leptocerus respectively (Reclaire 1936: 255; 1943:
181870):
Records. — ® Friesland: Terschelling, 21.vii.1954, W.
H. Gravestein (ZMA, P); Ibid., Oosterend, 24.viii.1962 &
3.ix.1969, J. H. Woudstra; Ibid., Bosplaat, 5.viii.1975,J.H.
Woudstra; Ibid., Hoorn, 28.vi.1978, J. H. Woudstra. ® N-
Holland: Bakkum, 28.vi & 19.vii.1969, J. H. Woudstra;
Ibid., 12.vii.1969. 29.vi.1971 & 18.vii.1978, B. Aukema. ®
Limburg: Colmont, 1.vii.1967, 1 9, P. Poot.
236. Heterocordylus leptocerus (fig. 49)
References. — Reclaire (1932: 195; 1936: 255; 1943:
117; 1948: 49; 1951: 18).
Published records. - ® Overijssel: Deventer (Reclaire
1932). ® Gelderland: Arnhem, Beek, Ermelo, Wagenin-
gen (Reclaire 1932); Doetinchem (Reclaire 1936); Gar-
deren (Reclaire 1943). ® Limburg: Meerssen, Mook (Re-
claire 1932); Herkenbosch (Reclaire 1948); Maasniel
(Reclaire 1951).
Unpublished records. - ® Overijssel: Oldenzaal, vii, 2
x Q, A. J. E Fokker (RML). ® Limburg: Maastrict, St.
Fig. 50. Mecomma ambulans. Pietersberg, 7.vi.1985, B. Aukema.
Present status. - Recorded only once since 1946.
238. Heterocordylus tumidicornis
References. — Reclaire (1940: 112); Blöte (1950: 19).
AI Published records. - ® Limburg: Eijs (ZMA) (Reclaire
ANS 1940); Maastricht (RML, NMM) (Blôte 1950).
Va | Present status. - Not recorded since 1949: Maastricht,
È St. Pietersberg, 13.vii, Blöte et al. (RML, NMM).
() 239. Heterotoma planicornis Pallas
© Synonym: H. meriopterum auct. nec Scopoli.
References. - Tamanini (1962) showed that what was
formerly considered to be H. meriopterum Scopoli in fact
| is planicornis (Pallas) (described from Belgium), whereas
the true meriopterum occurs east of the line Venice-Graz-
| Vienna.
241. Blepharidopterus diaphanus
References. - Kerzhner (1977) transferred diaphanus
from Orthotylus to Blepharidopterus because of the great
similarity in & genitalia between this species and B.
: angulatus.
242. Orthotylus fuscescens
References. - Klomp & Teerink (1973: 332).
Published records. - © Gelderland: Hoge Veluwe
(Klomp & Teerink 1973).
Unpublished records. - © Overijssel: Diepenveen, 22-
23.vi.1967, 1 @, J. J. Meurer, at light (ZMA). © Gelder-
land: Schaarsbergen, vi.1969, H. Klomp et al. (CB, AU).
© Limburg: Lomm, 8.vi.1982, B. Aukema & J. H. Woud-
Fig. 51. Myrmecoris gracilis. stra.
56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
248. Orthotylus virens
References. - Arnoud (1958).
Published records. — © Limburg: Eygelshoven,
16.vii.1958, 1 &, Br. Arnoud.
Unpublished records. - © Overijssel: Ootmarsum,
2.vii.1970, 1 4, at light, B. Aukema.
258. Cyrtorhinus caricis
References. — Reclaire (1932: 191).
Published records. - ® Friesland: Terschelling (Re-
claire 1932).
Unpublished records. — ® Friesland: West-Terschel-
ling (Groene Strand), 28-30.viii.1983 & 25.viii.1984, J. H.
Woudstra. ® Drenthe: Wijster, IJsbaan, 6 & 8.ix.1975 &
12.ix.1982, B. Aukema; Norg, Norgerholt, 24.viii.1977,
10 & 24.viii.1978, B. Aukema. ® Gelderland: Tongeren,
16.viii.1983 & 24.vii.1985, K. den Bieman; Ibid,
8.viii.1985, B. Aukema & K. den Bieman.
260. Mecomma ambulans (fig. 50)
References. — Reclaire (1932: 191).
Published records. - ® Gelderland: Winterswijk (Re-
claire 1932).
Unpublished records. - ® Overijssel: Denekamp, 18,
19.vi & 17.vii.1960, W. H. Gravestein (ZMA); Ibid.,
18.vi.1960, R. H. Cobben; Ootmarsum, 19.vi.1960, R. H.
Cobben; Ibid., Springendal, 5.vii. 1970 & 24.vii.1974,J.H.
Woudstra. ® Gelderland: Epe, Zuuk, 29.vii.1978 & Eper-
holt, 5.vii.1980, B. Aukema. ® Utrecht: Leersum,
8.vii.1981, 1 6, K. den Bieman. ® Limburg: Epen, Onder-
ste Bos, 29.vii.1938, H. C. Blöte (RML).
261. Myrmecoris gracilis (fig. 51)
References. - Reclaire (1932: 157; 1936: 253; 1951:
12); Cobben (1958: 18).
Published records. - ® Gelderland: Nijmegen, Wolf-
heze (Reclaire 1932); Groesbeek (Reclaire 1936). ® N-
Brabant: Oirschot (Reclaire 1951). ® Limburg: Horn
(Cobben 1958).
Unpublished records. — ® Drenthe: Zuidlaren,
18.vi.1950, W. H. Gravestein (ZMA); Wijster, 3 &
8.vii.1977, B. Aukema. © Gelderland: Arnhem,
28.vi.1908, Bierman (ZMA); Nijmegen, 22-23.vii.1933,
A. M. Scholte (RML, ZMA); Hatertse Vennen, 5.vii.1979,
R. H. Cobben. ® N-Brabant: Best, 21.vi.1950, M. van den
Boorn (P). ® Limburg: Mook, 23.vii.1926, F. T. Valck
Lucassen (ZMA).
263. Lygus adspersus f. nov. spec. (fig. 52)
References. - Kerznner (1984: 35-37); Rieger (1987:
278-281).
Records. - ® Overijssel: Holten, 7.x.1974, B. Aukema.
© Gelderland: Wageningen, 3.ix.1950, R. H. Cobben
(Cobben 1958a: 32, as. L. gemellatus); Ibid., 18.x.1987 &
23.x.1988, B. Aukema; Heelsum, 1.ix.1954, R. H. Cobben;
Ibid., 22.viii.1985, B. Aukema; Emst, 23.viii.1974 &
Heerde, 23.viii.1974, B. Aukema; Winterwijk, 17.ix.1983,
H.Vallenduuk (ZMA); Nijmegen, 6.x1.1987 & 28.x.1988,
B. Aukema; Culemborg, 13.x.1988, J. H. Woudstra; Elst,
28.x.1988, B. Aukema. ® Utrecht: Remmerden, 1.x.1988,
Fig. 52
. Lygus adspersus.
Fig. 53
. Lygus gemellatus.
AUKEMA: Checklist Hemiptera-Heteroptera 57
B. Aukema. ® N-Holland: Zaandam, 21.x.1976, B. Au-
„= kema. © Zeeland: Kloetinge, 10.v.1944, F Burger (RML).
® Limburg: Spaubeek, St. Odilienberg, Vlodrop, Wieler,
24-30.1x.1948, H. C. Blöte et al. (RML); Maastricht, St.
Pietersberg, 20.ix.1949 & Cannerbos, 20.11.1950, H. C.
Blöte et al. (RML); Elsloo, 12-20.ix.1950, H. C. Blöte et
al. (RML); Roermond, 4.x.1952 (Cobben 1958a: 35, as L.
gemellatus), St. Odiliënberg, 9.viii.1953, Herkenbosch,
10.1x.1954 & Melick-Herkenbosch, 21.viii.1955, R. H.
Cobben; Geulle, 21.ix.1975, B. Aukema.
Present status. - Widespread, especially along the lar-
ger rivers on Artemisia vulgaris L. and A. campestris L.
ee —
#
264. Lygus gemellatus (fig. 53)
Present status. — Revision of the Dutch material of
Lygus showed gemellatus as being limited to the south-
east part of the country.
265. Lygus maritimus (fig. 54)
References. — Gravestein (1951a: vi; 1951b: xxxvii-
XXXVIII).
Present status. - Widespread along the coast and along
the large rivers.
266. Lygus pratensis (fig. 55)
Present status. - More generally distributed than ge-
mellatus, occurring also in the western and northern
Fig. 54. Lygus maritimus. parts of the country. Rare.
268. Lygus wagneri f. nov. spec.
Records. - © Drenthe: Steenbergen, 6.x.1957, 1 8
(ZMA); Norg, Norgerholt, 30.ix.1971, J. Krikken & Ph.
Pronk (RML); Ibid., 30.vi.1975, 5 & 7.vi.1976, 3.v, 21.vi
& 24.viii.1977, 31.v, 10 & 24.viii.1978 & 16.v.1979, B.
Aukema; Ibid., 6.ix.1983, J. van Tol (RML); Vries, Zeijen,
10 & 24.viii.1978 & 16.v.1979, B. Aukema; Norg, Huis ter
Heide, 5.ix.1983, Ph. Pronk (RML); Roden, 6.ix.1983,
Ph. Pronk (RML).
Present status. — Very local, known only from the
boreal region around Norg.
270. Orthops basalis (fig. 56)
References. - Meurer (1956: 59); Cobben (1958a: 36-
40).
Present status. - Widespread, common.
275. Pinalitus atomarius
Note. - Kerzhner (1988) transferred a number of spe-
cies, including atomarius (Meyer-Dür), cervinus (Her-
rich-Schäffer), rubricatus (Fallen) and viscicola (Puton),
from the genus Orthops Fieber to Pinalitus Kelton.
References. - Cobben (1958a: 41).
Published records. - ® N-Holland: Heemstede,
26.11.1953, 1 Q, J. J. Meurer (ZMA).
Present status. - Not recorded since 1953.
276. Pinalitus viscicola
References. — Reclaire (1936: 253); Arnoud (1958);
Fig. 55. Lygus pratensis. Cobben (1958a: 40-41); Cobben & Arnoud (1969).
58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Fig. 56. Orthops basalis. Fig. 57. Plesiocoris minor.
Fig. 58. Agnocoris reclairei. Fig. 59. Agnocoris rubicundus.
AUKEMA: Checklist Hemiptera-Heteroptera 59
Published records. - ® Limburg: Epen, Mechelen (Re-
claire 1936; Cobben 1958); Voerendaal (Arnoud 1958);
Benzenrade, Nijswiller, Heerlen (Cobben & Arnoud
1969).
Unpublished records. - ® Limburg: Geulle, 16.viii &
27.ix.1966, P.J. Brakman (RML); Ibid., 27.ix.1966, P. Poot
(ZMA); Camerig, 26.viii.1973, B. Aukema; Ibid.
7.x.1973, B. Aukema & J. H. Woudstra; Noorbeek, 30.viii
& 3.ix.1973, B. Aukema; Ibid., 10.ix.1973, B. Aukema &
J. H. Woudstra.
277. Plesiocoris minor (fig. 57)
References. - Gravestein (1952a: cxvi-cxvii).
Present status. - Restricted to the West Frisian Islands
Vlieland, Ameland, Terschelling (Friesland) and Texel
(N-Holland).
281. Lygocoris populi f. nov. spec.
Records. - ® N-Holland: 's Graveland, 4.vii.1943, 1 9,
A. Reclaire (ZMA); Bakkum, 12.vii.1969, 1 9, B. Au-
kema; Ibid., 14.viii.1981 & 29.viii.1982, J. H. Woudstra;
Amsterdam, Bos, 24.vi.1973 & 18.vii.1981, B. Aukema. ®
Gelderland: Ubbergen, Beek, 16.vii.1987, B. Aukema &
D.J. Hermes; Doorwerth, 27.vi & 9.vii.1988, B. Aukema.
Present status. - Rare, on Populus canescens (Ait.) Sm.
283. Lygocoris limbatus
Fig. 60. Polymerus vulneratus. a — Reclaire (1932: 170; 1936: 253; 1943:
114).
Published records. - ® Gelderland: Leuvenum (Re-
claire 1936). ® N-Brabant: Oirschot (Reclaire 1932). ®
Limburg: Broekhuizen (Reclaire 1943).
Unpublished record. — © Overijssel: Losser,
17.vii.1960, W. H. Gravestein (ZMA). ® N-Brabant:
Biesbos, 8.v.1958, R. H. Cobben.
287. Agnocoris reclairei (fig. 58)
References. - Gravestein (1951a: v).
Present status. - Widespread.
288. Agnocoris rubicundus (fig. 59)
References. - Cobben (1958a: 41).
Note. - All records in Reclaire (1932: 172; 1934: 56;
1936: 253; 1940: 111; 1943: 115) concern A. reclairei
(Cobben 1958a).
Published records. - ® Limburg: Roermond, 10.v.1953,
1 g; Stein, 23.viii.1955, 1 9, R. H. Cobben (Cobben
1958a).
Unpublished records. - ® Utrecht: Rhenen, Blauwe
Kamer, 19.v.1985, B. Aukema; Ibid., 29.viii.1985, J. H.
Woudstra. ® Gelderland: Oosterbeek, 16.viii.1955, R. H.
Cobben; Valburg, Loenen, 2.viii.1985, B. Aukema. © N-
Brabant: Best, 15.iv.1977, H. W. van der Wolf (ZMA). ®
Limburg: Asselt, 2.vii.1974, B. Aukema; Wieler, 4.vii.
1974, B. Aukema; Echt, 4.vii.1982, C.J. M. Berger (ZMA);
Maastricht, St. Pietersberg, 29.viii. 1984, B. Aukema.
292. Polymerus vulneratus (fig. 60)
References. - Reclaire (1932: 174; 1943: 115).
Fig. 61. Polymerus holosericeus. Present status. - Before 1957 widespread in the Dune
60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
region and on the West Frisian Islands, but recently
recorded only once: N-Holland, Heemskerk, 22.viii.1987,
2 Q at light, J. H. Woudstra.
293. Polymerus holosericeus (fig. 61)
References. - Reclaire (1932: 175; 1948: 47; 1951: 14-
15); Gravestein (1943: ix); Cobben (1958: 19).
Published records. - ® Overijssel: Denekamp (Re-
claire 1951). ® Gelderland: Winterswijk (Gravestein
1943). e Limburg: Epen (Reclaire 1932); Arcen (Grave-
stein 1943); Vlodrop (Reclaire 1948); St. Odiliénberg
(Reclaire 1951; Cobben 1958).
Present status. - Not recorded since 1955: St. Odilien-
berg, 13.vii, R. H. Cobben (CB, ZMA).
294. Polymerus nigrita (fig. 62)
References. — Gravestein (1949: xxiii-xxiv); Reclaire
(1951: 15); Cobben (1958: 19).
Published records. — ® Overijssel: Denekamp (Grave-
stein 1949; Reclaire 1951). ® Limburg: St. Odiliënberg
(Cobben 1958).
Unpublished records. - ® Drenthe: Mantinge, 26.vi, 3
& 24.vii.1975, 19.vi.1978, B. Aukema. ® Gelderland:
Winterswijk, 22.vi.1952, P. J. Brakman (RML); Wamel,
21.vi.1981, 1 4, K. den Bieman; Ubbergen, Tiengeboden,
22.vi.1986, 1 6, K. den Bieman. ® Utrecht: Leersum,
Broekhuizen, 22.vi.1971, 1 9, M. P. Peerdeman (ZMA).
© Limburg: Bergen, De Hamert, 19.vi.1966, P. Poot;
Wieler, 4.vii.1973, B. Aukema; Cottessen & Epen, Bo-
venste Bos, 17.viii.1974, B. Aukema & J. Beernink; Vlo-
drop, 26.vi.1973, 1 9, B. van Aartsen (ZMA).
296. Dichrooscytus intermedius (fig. 63)
References. - Gravestein (1949: xxiii); Reclaire (1951:
14); Cobben (1953: 190-194; 1958: 19).
Published records. - ® Overijssel: Vasse (Gravestein
1949; Reclaire 1951). ® Gelderland: Wageningen (Cob-
ben 1953, 1958).
Unpublished records. - ® Drenthe: Assen, 7.vii.1964,
1 6, €. A. Schulz (ZMA). ® Overijssel: Hengelo,
2.vii.1952, W. C. Boelens (RML, P); Ootmarsum, Sprin-
gendal, 4 & 5.vii.1970, B. Aukema & J. H. Woudstra. ®
Gelderland: Winterswijk, 21.vi.1952, W. H. Gravestein
(ZMA); Wekerom, 1.vii.1958, 1 6, R. H. Cobben; Garde-
ren, 19.vii.1978, J. H. Woudstra; Epe, Eperholt, 6 &
15.vii.1980, B. Aukema; Ruurlo, 22. vii.1980, B. Aukema;
Doorwerth, 27.vi & 9.vii.1988, B. Aukema; Bennekom,
4 vii.1988, K. den Bieman. ® Utrecht: Maarn, 9.vii.1969,
J. Beernink; Broekhuizen, 5.vii.1973, 1 9, M. P. Peerde-
man (ZMA); Veenendaal, 30.vi.1988, B. Aukema. ® Lim-
burg: Epen, 14.vii.1966, J. H. Woudstra, Arcen,
16.vii.1980, 1 6, B. van Aartsen (ZMA).
299. Hadrodemus m-flavum (fig. 64)
References. — Fokker (1891: 359, as Calocoris margi-
nellus F.); Reclaire (1932: 166, as Homodemus m-flavum;
1948: 46; 1950: 13).
Published records. - ® Limburg: Nuth (Fokker: 1891);
Geulle, Mechelen, Mook and Valkenburg (Reclaire 1932);
Wijlre (Reclaire 1948); Epen (Reclaire 1950).
Present status. - Not recorded since 1949: Geulle, 12-
16.vi.1949, P. van der Wiel (ZMA).
Fig. 62. Polymerus nigrita.
Fig. 63. Dichrooscytus intermedius.
AUKEMA:
>
Fig. 64. Hadrodemus m-flavum.
Fig. 65. Megacoelum beckeri.
Checklist Hemiptera-Heteroptera 61
300. Calocoris pilicornis
References. - Gravestein (1952a: cxvi).
Published records. - © Limburg: Rijkel, 1.vii.1951, W.
H. Gravestein, R. H. Cobben & P. J. Brakman (Gravestein
1952a).
Present status. — Not recorded since 1953: Rijkel,
7.1.1953, W. H. Gravestein & P. J. Brakman (RML, ZMA,
AU, P & W).
301. Calocoris schmidtii
References. — Gravestein (1963).
Published records. - © Limburg: Gronsveld,
25.vi.1962, 1 6, W. H. Gravestein (ZMA) (Gravestein
1963).
Unpublished records. - ® Gelderland: Winterswijk, 't
Woold, 20.vi.1975, 1 6, B.J. Lempke (ZMA). ® Limburg:
Gronsveld, 15.vi.1968, W. H. Gravestein (ZMA); Ibid.,
23.vi.1968, P. Poot; Ibid., 4.vi.1969 & 10.vi.1981, J. H.
Woudstra; Ibid., 16.vi.1979, B. Aukema.
302. Calocoris striatellus
Synonym: C. gwadripunctatus Villers (Kerzhner 1969).
303. Calocoris biclavatus
References. - Reclaire (1951: 13).
Published records. — ® Limburg: Vaals, 24.vi.1948, 1 6,
P. van der Wiel (ZMA) (Reclaire 1951).
Present status. - Not recorded after the single capture
of 1948.
305. Calocoris affinis
References. - Reclaire (1943: 114).
Published records. - © Limburg: Gulpen, 21.vii.1943,
1 8, J. Maessen (Reclaire 1943).
Unpublished records. - ® Limburg: Epen, 29.vii.1949,
W. H. Gravestein (ZMA); Ibid., 26-27.vii.1943, H. C.
Blöte et al. (RML); Ibid, 25-26 & 30-31.vii.1964,
18.vii.1965, J. H. Woudstra; Ibid, 3-4.vii.1971 &
22.viii.1987, B. Aukema; Ibid, Bovenste Bos, 17.viii. 1974,
B. Aukema; Ibid., Onderste Bos, 3-4. vii. 1942, W. Vervoort
(RML); Gulpen, 20. vii. 1956, C. de Jong (RML); Heer-
len, 26.vii.1943, 1 9, H. C. Blöte et al. (RML); Wittem,
Bissen, 4.vii.1942, 1 9, W. Vervoort (RML).
312. Megacoelum beckeri (fig. 65)
References. - Cobben (1951: 52); Gravestein (1952a:
CXIV-CXVI).
Present status. - Widespread, but rare.
315. Miridius quadrivirgatus
References. - Fokker (1885: 56); Brakman (1952: 129).
Published records. - ® Zeeland: Zierikzee (Fokker
1885); Nieuw & St. Joosland (Brakman 1952).
Present status. - Not recorded since 1955: Nieuw & St.
Joosland, 15.viii.1955, P. J. Brakman (P).
317. Phytocoris intricatus (fig. 66)
References. - Reclaire (1951: 12); Cobben (1953: 190;
1958: 18).
62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Note. - The record from Nunspeet (Reclaire 1932)
concerns P. pim (Reclaire 1951).
Published records. - ® Drenthe: Norg (Reclaire 1951).
© Gelderland: Wageningen (Cobben 1953, 1958). ®
Overijssel: Vasse (Reclaire 1951).
Unpublished records. — ® Drenthe: Wijster, Zuid-
wolde (B. Aukema). ® Overijssel: Ootmarsum (B. Au-
kema, J. H. Woudstra). ® Gelderland: Apeldoorn, Win-
terswijk (W. H. Gravestein, ZMA); Brummen, Garderen,
Nunspeet (J. H. Woudstra); Epe, Ruurlo, Doorwerth (B.
Aukema); Bennekom (K. den Bieman). ® N-Holland:
Naarden, Zaandam (J. H. Woudstra). ® Limburg: Swal-
men (H. C. Blöte et al, RML); Epen, Schin op Geul (J.
H. Woudstra).
524. Phytocoris singeri (fig. 67)
References. — Cobben (1950: 84, as P nowickyi Fieber;
1958: 17-18).
Published records. - ® Limburg: Herkenbosch, Poster-
holt, St. Odiliënberg, Roermond, Neer and Vlodrop
(Cobben 1950, 1958).
Unpublished records. — ® Limburg: St. Odiliënberg,
9.viii.1953, 1 6, R. H. Cobben (LUW); Nuth, 13.viii. 1969,
1 é, J. H. Woudstra, Herkenbosch, Meijnweg,
2á.viii. 1968, 1 6, A. W. P. Maassen (RML); Cottessen, 26-
30.viii. 1968 & 10.ix.1973, J. H. Woudstra, Ibid,
17.viii.1971 & 29.viii.1973, B. Aukema; Ibid, 17.viii. 1974,
B. Aukema & J. Beernink; Epen, 14.viii.1971, 1 4, B.
Aukema; Colmont, Wrakelberg, 2.ix.1982, 1 @, K. den
Bieman; Maastricht, St. Pietersberg, 11.viii.1983, 1 3, B.
Aukema.
328. Capsus pilifer f. nov. spec. (fig. 68)
Records. — Revision of the Dutch Capszs material
resulted in the following localities for €. pilifer: ® Fries-
land: Appelscha, Elsloo (FNM); Beetsterzwaag (ZMA)
® Drenthe: Echten (RML); Lhee, Mantinge, Norg, Zuid-
wolde, Wijster (AU); Dwingeloo, Zuidlaren (ZMA). ®
Overijssel: Haaksbergen (RML); Ootmarsum (W). ®
Gelderland: Berg & Dal (JA); Ede (AU); Velp (ZMA);
Winterswijk (HE). ® Utrecht: Baarn (ZMA). ® N-Hol-
land: Texel (W).
Present status. - Widespread, but rare. On Molinia
caerulea (L.) Moench.
329. Capsus wagneri (fig. 69)
References. - Gravestein (1952a: cxvii).
Present status. - Widespread, rare.
331. Capsodes gothicus
References. - Snellen van Vollenhoven (1878a: 186-
187); Reclaire (1932: 179).
Published records. - © ?-Holland: Unknown locality,
W. de Haan, 1 6 (RML) (Snellen van Vollenhoven 1878).
© Limburg: Epen, vi.1911, 1 6, 1 9, D. MacGillavry
(ZMA) (Reclaire 1932).
Present status. - Not recorded since 1911.
332. Capsodes sulcatus f. nov. spec.
Records. — ® Gelderland: Vorden, Kranenburg,
23.vi.1979, 1 brachypterous Q, B. J. Lempke (ZMA).
Fig. 66. Phytocoris intricatus.
Fig. 67. Phytocoris singers.
63
AUKEMA: Checklist Hemiptera-Heteroptera
Fig. 69. Capsus wagneri.
Fig. 68. Capsus pilifer.
virens.
Fig. 71. Stenodema
Fig. 70. Stenodema trispinosa.
64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
ie
0 | «
Fig. 72. Megaloceroea recticornis. Fig. 73. Trigonotylus caelestialium.
Fig. 74. Trigonotylus ruficornis. Fig. 75. Teratocoris saundersi.
AUKEMA:
342. Trigonotylus caelestialium. Scale: 1 mm.
334. Acetropis gimmerthalii
References. - Fokker (1885: 54).
Published records. - © Z-Holland : ’s-Gravenzande,
Staelduin, 1 9, Snellen van Vollenhoven (RML) (Fokker
1885).
Present status. - Not recorded since 1878.
336. Stenodema trispinosa (fig. 70)
References. - Gravestein (1951b: xxxviii-xxxix); Brak-
man (1952: 130); Woudstra (1953); Meurer (1963: 205).
Present status. - Widespread, mainly coastal.
Checklist Hemiptera-Heteroptera 65
339. Stenodema virens (fig. 71)
References. — Reclaire (1932: 180-181; 1936: 254;
1943: 116; 1948: 47; 1951: 16).
Present status. - Until 1946 widespread, but since then
only two records: Friesland, Terschelling, Bosplaat,
12.viii.1964, J. Beernink; Gelderland, Emst, 23.viii.1973,
B. Aukema.
341. Megaloceraea recticornis (fig. 72)
References. - Blöte (1950: 19); Cobben (1953: 194);
Brakman (1960: 22).
Present status. - Before 1950 hardly any records out-
side the southern part of Limburg. At present however
widespread.
342. Trigonotylus caelestialium f. nov.
spec. (fig. 73)
345. Trigonotylus ruficornis (fig. 74)
References. - Bozdechovä (1973).
Present status. - Revision of Dutch Trigonotylus ruft-
cornis revealed that both 7: caelestialium and T. ruficor-
nis are common species in the Netherlands.
347. Teratocoris paludum
References. — Cobben (1960: 206-207).
Published records. - © Gelderland: Wageningen (Cob-
ben 1960).
Unpublished records. - ® Drenthe: Wijster, IJsbaan,
6.1x.1975, 10 & 11.vi.1976 & 12.ix.1982, B. Aukema; Ibid.,
26.viii.1976, B. Aukema & J. H. Woudstra; ® Gelderland:
Elst, Elden, 20.vii.1947, 1 9, A. Reclaire (ZMA); Tonge-
ren, 8.111.1985, B. Aukema & K. den Bieman; Ibid,
19.ix.1985, K. den Bieman. ® Utrecht: Veenendaalse Hel,
7&11.viii.1981, G. J. Rozeboom & R. H. Cobben (LUW).
348. Teratocoris saundersi (fig. 75)
References. — Fokker (1891: 359); Reclaire (1940:
111); Gravestein (1952a: cxvii-cxviii, as T. lineatus Wag-
ner); Cobben (1958: 19); Brakman (1960: 22, as T. line-
atus).
Published records. - © N-Holland: Wieringen (Re-
claire 1940); Texel, Slufter (Gravestein 1952a). ® Z-
Holland: Oostvoorne (Cobben 1958). ® Zeeland: Zierik-
zee (Fokker 1891); Hoek (Brakman 1960).
Unpublished records. - ® Friesland: Terschelling (W.
H. Gravestein, J. H. Woudstra, B. Aukema); Ameland (B.
Aukema). ® N-Holland: Naarden (J.J. Wieringa, M). ®
Z-Holland: Oostvoorne (ZMA), Goeree (K. J. Huisman,
RML). ® Zeeland: Walcheren (H. C. Blöte, RML); West-
kapelle, Ellewoutsdijk (B. Aukema).
Present status. - Widespread, mainly coastal.
351. Isometopus intrusus (fig. 76)
References. — Fokker (1886: 299, as I. alienus Fieber;
1891: 360); Reclaire (1940: 113; 1943: 120).
Published records. - ® Gelderland: Winterswijk (Fok-
ker 1891). © N-Brabant : Waalwijk (Reclaire 1940). ©
Limburg: Valkenburg (Fokker 1886); Arcen (Reclaire
1943).
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
66
Fig. 77. Stalia boops.
Fig. 76. Isometopus intrusus.
Fig. 79. Nabis pseudoferus.
Fig. 78. Nabis brevis.
AUKEMA: Checklist Hemiptera-Heteroptera 67
Fig. 80. Temnostethus longirostris.
Fig. 81. Elatophilus nigricornis.
Unpublished records. - ® Gelderland: Wageningen, vii
& viii.1960, vii.1961 & viii.1962, R. H. Cobben (CB, LUW,
ZMA, AU). ® Utrecht: Hollandse Rading, 8.viii.1956, 1
Q, J. H. Woudstra.
352. Prostemma guttula
References. — Snellen van Vollenhoven (1878a: 305-
306); Reclaire (1932: 141; 1936: 251); Brakman (1952:
129); Cobben (1958: 16); Péricart (1987: 50).
Published records. - ® Limburg: Maastricht (Reclaire
1932; 1936); Echt (Cobben 1958). ® Zeeland: Walcheren
(Snellen van Vollenhoven 1878); Domburg (Brakman
1952).
Unpublished records. — ® Limburg: Maastricht, St.
Pietersberg, 20.viii.1983, 1 9, ex larva, B. Aukema. ®
Zeeland: Domburg, 1957.viii.13, P. J. Brakman (RML).
353-364. Nabini
Note: Kerzhner (in litt., 1989) after analysis of the
world fauna now considers Aptus, Anaptus and Stalia as
subgenera of Himacerus, and Limnonabis, Dolichonabis
and Nabicula as subgenera of Nabis. However, since the
recent work of Péricart (1987) on this subject probably
will remain the standard in Western Europa for many
years, I prefer to maintain his classification.
356. Stalia boops (fig. 77)
References. - Reclaire (1936: 251); Leston (1952: 86);
Gravestein (1955); Cobben (1958a: 17); Brakman (1960:
22); Péricart (1987: 93).
Present status. - Widespread, but rare.
360. Nabis brevis (fig. 78)
Note. - Most older identifications are unreliable (Re-
claire 1951)
References. — Reclaire (1936: 251; 1948: 44; 1951: 11);
Cobben (1958: 16); Péricart (1987: 144-147).
Published records. - ® Gelderland: Hoog-Buurlo (Re-
claire 1936); Nunspeet (Reclaire 1936, 1951); Hulshorst
(Reclaire 1948); Lunteren (Reclaire 1951); Ede (Cobben
1958). ® Limburg: Haelen (Reclaire 1951).
Unpublished records. — ® Overijssel: Ootmarsum,
Vasse (J. H. Woudstra). ® Gelderland: Ede (R. H. Cob-
ben); Emst, Gortel, Heerde, Wezep (B. Aukema); Vier-
houten (B. Aukema; J. Beernink); Epe (B. Aukema; Ph.
van Hooven, RML; C. Scheffer, RML); Putten (C. de
Jong, RML); Eerbeek (H. Boschma, RML); Ugchelen (P.
J. Hartevelt & Ph. Pronk, RML); Velp (W. H. Gravestein,
P). © N-Brabant: Herpen (S. v. Heijnsbergen, W); Best,
Strijbeek (B. Aukema; J. H. Woudstra); Best, Nijnsel (H.
W. van der Wolf, ZMA). ® Zeeland: Axel (C. van Heij-
ningen, RML); Hulst (Ph. Pronk, RML). ® Limburg:
Maastricht (B. Aukema).
363. Nabis pseudoferus (fig. 79)
References. - Gravestein (1956); Péricart (1987: 137).
Present status. - Widespread, rare.
366. Temnostethus longirostris f. nov. spec.
(fig. 80)
Records. - ® Overijssel: Steenwijk, 4.viii.1987, B. Au-
68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
kema; Zwolle, Schelle, 21.vii.1987. ® Flevoland: Lelystad,
Oostvaardersplassen, 22.vi.1988, B. Aukema. ® Gelder-
land: Wageningen, 10-20.vii.1987, B. Aukema; Millingen,
16.vii.1987, B. Aukema; Ooij, Groenlanden, 16.vii.1987,
B. Aukema; Heelsum, 26.vi.1988, B. Aukema; Door-
werth, 27.vi.1988, B. Aukema; Doornenburg, 2.vii.1988,
B. Aukema. ® Utrecht: Rhenen, Blauwe Kamer,
20.vii. 1987, B. Aukema. © N-Holland: Zaandam, 19.vii-
19.viii.1987, 21.vii & 17.viii.1988, J. H. Woudstra (W,
ZMA); Zaandijk, 22.vii.1987, J. H. Woudstra; Kromme-
nie, 28.viii.1988, B. Aukema. ® Limburg: Swalmen,
1.vii.1951, P. van der Wiel (ZMA).
Present status. - Widespread, on trunks of Salix and
Populus.
368. Elatophilus nigricornis (fig. 81)
References. - Cobben & Arnoud (1969: 8).
Published records. — ® Friesland: Beetsterzwaag (Cob-
ben & Arnoud 1969). ® Overijssel: Lutterzand, Ootmar-
sum (Cobben & Arnoud 1969).
Unpublished records. — ® Drenthe: Spier, Kibbelhoek,
10-14.vi.1977, B. Aukema. © Gelderland: Heerde,
20.vi.1971, 1 Q, B. Aukema. ® Utrecht: Zeist, 1á.v. 1969,
J. Burger (PDW). ® N-Brabant: Drunen, 4.vi.1983, A. P.
J. A. Teunissen (ZMA). ® Limburg: Lomm, Ravenven-
nen, 6.vi.1982, B. Aukema.
369. Anthocoris amplicollis
References. - Aukema (1976).
Published records. — © Limburg: Bunde (Aukema
1976).
Unpublished records. — ® Utrecht: Amerongen,
28.ix.1986 & 30.viii.1987, K. den Bieman; Ibid.
15.viii.1987, B. Aukema; Zeist 16.viii.1987, 1 6, K. den
Bieman.
370. Anthocoris butleri (fig. 82)
References. - Cobben & Arnoud (1969: 7-8).
Present status. — Locally common on Buxus in Gelder-
land, Utrecht, N-Holland and Limburg.
374. Anthocoris minki minki (fig. 83)
References. - Most older records concern the Ash
(Fraxinus exselsior L.) inhabiting A. simulans, establis-
hed by Péricart (1972: 139-141) as a good species.
Present status. - The true Poplar (Populus) inhabiting
A. minki is known from a few localities only, but it is
undoubtedly a widespread species in the Netherlands.
377. Anthocoris pilosus (fig. 84)
Synonym: A. sibiricus auct nec Reuter (Elov & Kerzhner
1977)
References. - Reclaire (1932: 149; 1936: 252); Blöte
(1950: 19, 1954: 83); Cobben (1953: 190); Péricart (1972:
143-145, as szbiricus).
Note. - The specimen recorded as sibiricus from
Oisterwijk (Reclaire 1932) was not traceable.
Published records. - ® Limburg: Maastricht (Reclaire
1930; Blöte 1950, 1954), Meerssen (Reclaire 1932); Epen,
Eygelshoven (Reclaire 1936); Roermond, Echt, Maasniel,
Swalmen (Cobben 1953).
Fig. 82. Anthocoris butleri.
Fig. 83. Anthocoris minki minki.
AUKEMA:
Fig. 84. Anthocoris pilosus.
Fig. 85. Anthocoris simulans.
Checklist Hemiptera-Heteroptera 69
Present status. - Recorded only once since 1951: Maas-
tricht, 24.viii.1965, P. Poot.
379. Anthocoris simulans (fig. 85)
Note. - see under A. minki.
Present status. - Widespread and common.
380. Anthocoris visci
References. - Cobben & Arnoud (1969: 5-7).
Published records. - ® Limburg: Benzenrade, Holset,
Mechelen, Nijswiller, Putberg, Geulle (Cobben & Arnoud
1969).
Unpublished records. - ® Limburg: Geulle, 27.ix.1966,
P. Poot (ZMA, P); Noorbeek, 30.viii, 3 & 8.1x.1973, B.
Aukema. Ibid., 7.ix.1973, B. Aukema.
382. Acompocoris alpinus (fig. 86)
References. — Fokker (1885: 75).
Note. - The specimen recorded from Vorden (Fokker
1885) belongs to Tetraphleps bicuspis (Herrich-Schäf-
fer).
Published records. - ® Utrecht: Driebergen (Fokker
1885).
Unpublished records. - ® Friesland: West-Terschel-
ling J. H. Woudstra); Appelscha (J. H. Woudstra). ®
Drenthe: Wijster, Spier (B. Aukema). ® Gelderland: Epe,
Gortel, Heerde (B. Aukema); Garderen, Hulshorst (J. H.
Woudstra); Planken Wambuis (K. den Bieman); Nun-
speet (R. T. Simon Thomas, ZMA). ® Utrecht: Baarn
(RML); Hollandse Rading (J. H. Woudstra). © N-Hol-
land: Santpoort (B. Aukema; J. H. Woudstra); Naar-den,
Vogelenzang, Zaandam (J. H. Woudstra). ® N-Brabant:
Strijbeek (J. H. Woudstra), Zundert (J. H. Woudstra; P.
Aukema, AU). ® Limburg: Brunssum (K. den Bieman);
Epen (B. Aukema); Terziet (P. Poot); Vijlen (J. H. Woud-
Stra).
Present status. - A. alpinus was collected twice before
1885 (Driebergen, Baarn), whereas all other records are
from 1967 onwards.
385. Orius laticollis (fig. 87)
References. - Cobben (1958: 17).
Published records. - © Limburg: Roermond (Cobben
1958).
Unpublished records. — © Friesland: Terschelling,
Midsland (J. H. Woudstra). ® Drenthe: Norg (B. Au-
kema). ® Flevoland: Lelystad (H. Vallenduuk). ® Gelder-
land: Welsum (B. Aukema). ® Utrecht: Breukelen (C. van
Heijningen, RML); Kamerik, Remmerden (B. Aukema).
© N-Holland: Amsterdam, Krommenie (B. Aukema);
Ilpendam, Koog aan de Zaan, Westzaan (J. H. Woudstra).
Zaandam (J. H. Woudstra; J. Beernink); Heemskerk (J.
Beernink). ® Limburg: Cottessen, Epen (B. Aukema);
Maastricht (P. Poot); Mechelen (J. H. Woudstra).
Present status. - Widespread, but certainly not as rare
as the present records may suggest. Much older material,
identified as O. minutus still has to be checked for the
presence of this species and the next one.
388. Orius vicinus
References. - Cobben (1958: 17).
70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Fig. 86. Acompocoris alpinus. Fig. 87. Ortus laticollis.
Fig. 88. Xylocoris formicetorum. Fig. 89. Brachysteles parvicornis.
AUKEMA: Checklist Hemiptera-Heteroptera 71
Published records. — ® Gelderland: Wageningen (Cob-
ben 1958).
Unpublished records. — © Gelderland: Ugchelen,
18.ix.1960, 1 g, P. J. Hartevelt & Ph. Pronk (RML);
Zutphen, ix.1963, P. H. van Doesburg (RML); Ben-
nekom, 25 & 30.viii. 1986 & 7.ix.1987, K. den Bieman;
Wageningen, 4.ix. 1988, B. Aukema. © N-Holland: Am-
sterdam, 19.viii.1974 & 11.v.1977, B. Aukema; Ibid., 13 &
23.ix.1985, J. H. Woudstra. © Lim- burg: Epen, 3-
4.vii.1971, 1 À, & 11.ix.1980, 1 6, B. Aukema.
Present status. — Probably widespread, but still very
few times recorded with certainty.
392. Xylocoris formicetorum (fig. 88)
References. — Reclaire (1932: 153-154).
Published records. - € Gelderland: Otterloo (Reclaire
1932).
© Utrecht: Baarn (Reclaire 1932). © N-Holland: Hil-
versum (Reclaire 1932).
Unpublished records. — © Overijssel: Ommen,
viii.1986 & x.1987, H. Vallenduuk. ® Gelderland: Garde-
ren, 8.1.1983, H. Vallenduuk (ZMA); Nunspeet, iv.1987
& Kootwijk, vi.1987, H. Vallenduuk. © Utrecht: Doorn,
28.v.1984, H. Vallenduuk (ZMA); Driebergen, vii. 1987,
H. Vallenduuk. ® Z-Holland: Den Haag, 20.iii.1983, H.
Vallenduuk (ZMA). ® Zeeland: Haamstede, ix.1987, H.
Vallenduuk. ® Limburg: Horst, vii & x.1987, H. Vallen-
duuk.
Fig. 90. Oeciacus hirundinis.
393. Brachysteles parvicornis (fig. 89)
References. - Reclaire (1932: 154; 1934: 53; 1951: 12);
Cobben (1958: 17).
Published records. - © Gelderland: Leuvenum (Re-
claire 1934). ® Utrecht: Soest (Reclaire 1932). © N-
Holland: Hilversum (Reclaire 1932); Texel, De Koog
(Reclaire 1951). ® Z-Holland: Meijendel (Reclaire
1932). © Zeeland: Oostkapelle (Reclaire 1951). ® Lim-
burg: Roermond (Reclaire 1951); Boukoul, Roermond,
Bergen (Cobben 1958).
Unpublished records. — © Gelderland: Otterlo,
10.viii.1954, 1 9, W. H. Gravestein (ZMA). ® N-Hol-
land: Hilversum, 19.v.1947, A. Reclaire (P); Bakkum,
5.vi.1975, 1 @, J. Beernink. © Zeeland: Vrouwenpolder,
1.v.1949 & 29.vii.1954, P. J. Brakman (RML); Goes,
19.viii.1953, A. Post (ZMA); Oostkapelle, 3.x.1954, P. J.
Brakman (P).
Present status. - Formerly a widespread species, but
recorded only once since 1954.
394. Cardiastethus fasciiventris
References. - Reclaire (1936: 252).
Published records. — © Gelderland: Terborg,
3.viii.1935, 1 9, P. van der Wiel (ZMA) (Reclaire 1936).
Present status. - Not recorded since 1937: Terborg, 1-
4.viii, P. van der Wiel (ZMA).
396. Scoloposcelis pulchella f. nov. spec.
Records. - ® N-Holland: ’s Graveland, 15.vii.1977, 1
Fig. 91. Rhynocoris annulatus. Q at light, S. van Heijnsbergen (W).
72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
397. Oeciacus hirundinis (fig. 90)
References. — Fokker (1885: 78); Reclaire (1932: 147).
Published records. —: ® Gelderland: Arnhem (Reclaire
1932). ® Limburg: Venlo (Fokker 1885).
Unpublished records. — © Overijssel: Dalfsen,
11.vi.1977, ABW (AU); Zwolle. 3.v.1977, ABW (AU). ®
Gelderland: Overasselt, 30.ix.1947, F. Smit (ZMA, P);
Echteld, IJzendoorn, 22.v.1974 (PDW); Arnhem,
11.vi.1977, ABW (AU). ® Utrecht: Kockengen,
9.vii.1973, W. J. Klok (ZMA, AU). ® Z-Holland: Dord-
recht, 3.v.1977, ABW (AU). ® N-Brabant: Made,
4.vi.1970 (PDW); Helmond, 15.v.1974 (PDW); Andel,
14.vi.1977, ABW (AU). © Zeeland: Philippine,
14.vi.1974, ABW (AU); Goes, Kattendijke, 29.v.1975, B.
Bosman (PDW) Wemeldinge, 10.vi.1977, ABW (AU);
Yerseke, 10.vi.1977, ABW (AU).
399. Cimex columbarius
References. - Fokker (1885: 78); Reclaire (1932: 146;
1936: 251); Péricart (1972: 294-296).
Published records. - ® Groningen: Scheemda (Reclaire
1936; Péricart, 1972). ® Gelderland: Nijkerk (Reclaire
1936). ® Utrecht: Utrecht (Reclaire 1932). ® Z-Holland:
Giessendam (Fokker 1885).
Unpublished records. - © Groningen: Scheemda,
12.xii.1938 (P). ® N-Holland: Aalsmeer, 14.vi.1924, 2 Q,
D. MacGillavry (ZMA).
Present status. - Not recorded since 1938.
400. Cimex dissimilis.
References. - Fokker (1885: 78, as C. pipistrellae Ko-
lenati); Reclaire (1932: 146, as pipistrelli; 1936: 251, as
pipistrelli and dissimilis; 1943: 112, as pipistrelli); Péri-
cart (1972: 296-298, as pipistrelli and dissimili).
Published records. - ® N-Holland: Amsterdam, Aals-
meer, Haarlem (Reclaire 1936 1943).
Unpublished records. - ® Utrecht: Driebergen, viii, J.
Oudemans (RML, Museum Helsinki) (Fokker 1885 and
Péricart 1972, as pipistrelli); Groenekan, ix.1936, 2 9, H.
Vink (ZMA).
Present status. — Not recorded since 1940: Haarlem,
De Hout, 19.ix, L. Bels (ZMA).
401. Cimex pipistrelli
References. — Fokker (1885: 78); Péricart (1972: 296-
297).
Note. - With the exception of Zierikzee (Fokker 1885)
all published records of C. pipistrelli thusfar (Fokker
1885; Reclaire 1936, 1943; Péricart 1972) concern speci-
mens of C. dissimilis.
Published records. — ® Zeeland: Zierikzee (Fokker
1985).
Unpublished records. — ® Friesland: Berlikum, 28 &
30.vi.1967, P. E van Heerdt (ZMA); Tjerkwerd,
14.11.1977, A. M. Voûte et al. (ZMA); Ibid., 29.11.1977, B.
Aukema.
402. Empicoris baerensprungi
References. — Reclaire (1932: 139).
Published records. - ® Utrecht: Baarn (Reclaire 1932).
Fig. 92. Coranus subapterus.
Fig. 93. Coranus woodroffei.
AUKEMA: Checklist Hemiptera-Heteroptera 15)
® Gelderland: Leuvenum (Reclaire ,1932).
Unpublished records. - ® Gelderland: Hall, 5.vii.1987,
1 g, B. Aukema.
405. Pygolampis bidentata
References. — Fokker (1886: 298); Reclaire (1932:
139); Arnoud (1958); Cobben (1958: 16).
Published records. — ® Gelderland: Vorden, ix, 1 larva,
Groll (Fokker 1886). © N-Brabant: Deurne, 17.vi.1914,
1 6, D. MacGillavry (Reclaire 1932). © Limburg: Linne,
25.v.1944, 1 larva; Herkenbosch, 11.viii.1947, young lar-
vae, R. H. Cobben (Cobben 1958); Colmont, 19.vi.1958,
larva, Br. Arnoud (Arnoud 1958).
Unpublished records. - © N-Brabant: vicinity of Eind-
hoven, 1 8, C. Berger (P).
Present status. - Not recorded since 1958.
407. Rhynocoris annulatus (fig. 91)
References. - Snellen van Vollenhoven (1878a: 310, as
Harpactor annulatus); Fokker (1884: 129; 1899: 35); Re-
claire (1932: 140, as Rhinocoris iracundus (Poda); 1951:
10).
Published records. - ® Drenthe: Dwingeloo (Reclaire
(1951). ® N-Brabant: Ulvenhout (Snellen van Vollenho-
ven 1878); Eindhoven (Reclaire 1950). ® Limburg:
Gronsveld, Venlo (Fokker 1899); Valkenburg (Reclaire
1932).
Unpublished records. - © Drenthe: Dwingeloo,
11.vi.1967, 1 6 (ZMA); Wijster, 28.v.1978, 1 & 8.vi.1979,
28.v, 6 & 10.vi.1980, 6-7.viii.1984, in window traps, B.
Aukema (AU, W). © N-Brabant: Ginneken, 9.vi.1878, 1
®, E.J. M. Heylaerts (ZMA); Oisterwijk, 17.v. 1894, 1 6
(RML). © Limburg: Meerlo-Wanssum, Geijsteren,
20.v.1950, M. van den Boorn (P).
Fig. 94. Aneurus laevis.
408. Coranus subapterus (fig. 92)
409. Coranus woodroffei f. nov. spec.(fig. 93)
References. — Putshkov (1982).
Revision of material identified as C. subapterus reve-
aled both subapterus and woodroffei as widespread and
common species in the Netherlands.
410. Phymata crassipes
References. - Fokker (1899: 34).
Published records. - © Gelderland: Groesbeek,
27.vii.1890, 1 specimen, Groll (RML) (Fokker 1899).
Present status. - Not recorded since 1890.
412. Aradus corticalis
References. - Fokker (1899: 35); Cobben (1987).
Published records. - © Gelderland: Winterswijk,
xi.1910, 1 Q and exuvium, D. L. Uyttenboogaart (ZMA)
(Cobben 1987). ® Z-Holland: Den Haag, vi.1894, Everts
(RML) (Fokker 1899).
Present status. - Not recorded since 1910.
414. Aradus signaticornis
References. - Cobben (1987).
Fig. 95. Piesma quadratum. Published records. - © Gelderland: Wageningen,
74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
15.v.1985, 1 6, R. H. Cobben (Cobben 1987).
Unpublished records. - © N-Brabant: Helenaveen, 10-
30.viii.1987, K. Alders (ZMA); Ibid., 9.ix.1987, B. Au-
kema.
415. Aneurus avenius
References. - Aukema (1976b).
Published records. - © Limburg: Epen, Terziet, Val-
kenburg, Vijlen (Aukema 1976b).
Unpublished records. - © Drenthe: Mantinge,
31.v.1981, 1 9, Y. Jongema (AU). ® Limburg: Vijlen,
9.iii.1965, 1 4, Br. Arnoud.
416. Aneurus laevis (fig. 94)
References. - Aukema (1976b).
Present status. - Widespread.
418. Piesma quadratum (fig. 95)
References. - Reclaire (1932: 125; 1936: 249; 1940:
108; 1951:9); Woudstra (1959).
Note. - The specimens from Texel (Reclaire 1951)
belong to P. salsolae (Gravestein 1958).
Published records. - ® N-Holland: Zaandam (Woud-
stra 1959). ® Zeeland: Zierikzee, Walcheren (Reclaire
1932); Middelburg (Reclaire 1936); Rilland-Bath (Re-
claire 1940); Nieuw en St. Joosland, Vrouwenpolder (Re-
claire 1951).
Unpublished records. - ® Drenthe: Wijster, 22.i &
6.vii.1976, B. Aukema; Ibid., 31.viii.1977, J. H. Woudstra.
© N-Holland: Camperduin, 15.ix.1962 & 19.ix.1967, J.
Beernink; Ibid., 2.ix.1967, J. H. Woudstra; Zaandam,
10.ix.1960, J. Beernink. ® Z-Holland: Oost-Voorne,
9.vi.1969 & 19.vi.1970 (ZMA). ® Zeeland: Vrouwenpol-
der, 8.vii.1954 & Nieuw- en St. Joosland, 28.ii.1965, P. J.
Brakman (P); Krabbendijk, 14.viii.1983, 1 g, K. den
Bieman; Vlissingen, 21 & 27-28.vii.1986, B. Aukema. ®
N-Brabant: Breda, 27.vi.1964, H. J. van der Krift (ZMA);
Bergen op Zoom, 19.ix.1983, B. van Aartsen (ZMA). ®
Limburg: Beegden, 29.v.1980, K. den Bieman.
419. Piesma salsolae
References. — Gravestein (1959).
Published records. - © N-Holland: Texel, Slufter,
26.viii. 1946 & Ibid., De Koog, 26.viii.1946 & 29.vii. 1947,
W. H. Gravestein (ZMA).
Present status. - Only recorded from the West Frisian
Island of Texel, where it was also caught by J. H. Woud-
stra (6.1x.1972) and B. Aukema (29.viii.1974).
424. Berytinus montivagus
References. — Fokker (1883: 248; 1885: 51; 1891: 357).
Note. - The specimen recorded by Fokker (1883) from
Utrecht is not traceable and therefore ommitted from the
list. The record from Limburg (Heerderberg) in Blöte
(1954) concerns B. minor.
Published records. - ® Zeeland: Zierikzee (Fokker
1885; 1891).
Unpublished records. — ® Zeeland: Vlissingen, 15, 16
& 21.vii.1986, B. Aukema.
428. Metatropis rufescens
References. - Loof (1955); Cobben (19562).
Published records. — ® Limburg: Valkenburg (Loof
1955); Swalmen (Cobben 1956a).
Unpublished records. — ® Overijssel: Ootmarsum, 7-
9.vii.1972, J. H. Woudstra & B. Aukema. @ Limburg:
Bunde, 8.vi.1981, 1 4, K. den Bieman.
429. Lygaeus equestris
References. — Snellen van Vollenhoven (1878a: 91-92);
Reclaire (1936: 246; 1951: 246).
Note. - The record from Amsterdam (Reclaire 1951)
most likely concerns an imported specimen.
Published records. — ® Friesland: locality unknown
(Snellen van Vollenhoven 1878). ® Limburg: Maastricht
(Reclaire 1936).
Present status. - Not recorded since 1935: Maastricht,
3.vii, Maessen (NMM).
430. Spilostethus saxatilis
References. - Fokker (1884: 113, as Lygaeus saxatilis).
Published records. - ® Limburg: Maastricht, A. H.
Maurissen (RML) (Fokker 1884).
Present status. - Not recorded since 1884.
435. Ortholomus punctipennis
References. — Brakman (1952: 130, as Nysius puncti-
pennis); Gravestein (1952a: cxviii); Cobben (1958: 16);
Aukema & Woudstra (1985).
Published records. - © Zeeland: Domburg (Brakman
1952; Gravestein 1952). ® Z-Holland: Voorne, ’s-Gra-
venzande (Cobben 1958). ® N-Holland: Egmond aan
Zee (Aukema & Woudstra 1985).
Unpublished records. - ® N-Holland: Egmond aan
Zee, 14.viii.1985, J. H. Woudstra; IJmuiden, 14.viii.1986,
B. Aukema; Ibid., 21.viii.1986, J. H. Woudstra.
436. Orsillus depressus
References. - Aukema (19882).
Published records. - ® Gelderland: Nijmegen, Wage-
ningen (Aukema 1989a). ® N-Holland: Naarden (Au-
kema 1989a).
Present status. — Collected since 1985 on Chamae-
cyparis lawsoniana (Murr) Parl., apparently spreading.
Introduced with seeds of Thuja occidentalis L. in 1966
(PDW).
437. Kleidocerys privignus f. nov. spec.
(fig. 96)
Records: ® Zeeland: Domburg, Westhoven,
28.vi.1983, B. Aukema; Oostkapelle, 1.vii.1983 &
18.vii.1986, B. Aukema. ® N-Brabant: Eindhoven, Bokt,
24.iv & 12.xii. 1982, 15.v.1985, H. W. van der Wolf; Ibid.,
16.v.1985, B. Aukema & H. W. van der Wolf; Ibid. & De
Bochten, 30.xii.1982, B. Aukema, H. W. van der Wolf &
J. H. Woudstra; Nederwetten, 2.1.1983, H. W. van der
Wolf; Haarsteeg, 27.11.1983, Oudheusden, 5.111.1983 &
Capelle, 13.11.1983, A. P. J. A. Teunissen; Soerendonk,
20.1.1983 & Son, 12.x1.1987, H. W. van der Wolf. ®
Limburg: Swalmen, Wieler, 29.iv.1983, B. Aukema.
Fig. 96. Kleidocerys privignus.
Fig. 97. Cymus melanocephalus.
Checklist Hemiptera-Heteroptera 75
Present status. - Widespread in the southern provinces
on alder (Alnus glutinosa (L.) Gaertner), especially near
and above water. Rare.
439. Kleidocerys truncatulus ericae f. nov. spec.
Note. - Slater (1964) erroneously lists K. truncatulus
ericae for the Netherlands due to misinterpretation of
Reclaire (1932).
Records. - ® N-Holland: Texel, 't Mientje, 6.ix.1972,
J. H. Woudstra; Ibid., 29.viii.1974, B. Aukema.
Present status. — Collected on Calluna vulgaris (L.)
Hull on one locality only. Despite some efforts not found
there again since 1974.
440. Cymus aurescens
Synonym: C. obliguus Horvath (Josifov & Kerzhner
1978).
443. Cymus melanocephalus (fig. 97)
References. - Cobben (1948: 50); Reclaire (1951: 6).
Present status. — After its discovery in 1947 C. mela-
nocephalus has appeared to be widespread and locally
common.
445. Geocoris ater
References. — Snellen van Vollenhoven (1878a: 134-
135).
Published records. - ® Utrecht: Driebergen, vii, 1 9,
G. A. Six (RML).
Present status. - Not recorded since 1878.
447. Geocoris megacephalus
References. — Snellen van Vollenhoven (1878a: 135, as
Ophthalmicus pallidipennis); Reclaire (1932: 98-99).
Published records. - ® Zeeland: Vlissingen (Snellen
van Vollenhoven 1878); Cadzand, Domburg (Reclaire
1932).
Present status. - Not recorded since 1902: Cadzand, vii,
A. J. F. Fokker (RML).
448. Metopoplax ditomoides
References. - Reclaire (1951).
Published records. - ® Limburg: Gronsveld,
21.vi.1948, 1 6, P. van der Wiel (ZMA) (Reclaire 1951).
Present status. - Not recorded since 1948.
449. Oxycarenus modestus (fig. 98)
References. - Cobben (1951: 52-53); Aukema (1976a)
Published records. - © Limburg: Melick-Herkenbosch
(Cobben 1951); Wieler, Jabeek (Aukema 1976a).
Unpublished records. — © Gelderland: Dreumel,
11.iv.1981, 1 9, K. den Bieman; Lienden, 27.iii.1983, 1 4,
R. H. Cobben. © N-Brabant: Eindhoven, Bokt, 30.x.1977,
1 6, H. W. van der Wolf; Ibid., 30.xii. 1982; Vlijmen,
15.v.1982, A. P. J. A. Teunissen (ZMA); Ibid., 5.vii. 1982,
C.J. M. Berger (ZMA, P); Haarsteeg, 13.iii.1983, A. P. J.
A. Theunissen (ZMA); Best, 16.v.1985, B. Aukema. ®
Limburg: Wieler, l.viii.1980 & 29.iv.1983, B. Aukema;
Epen, 8.vi.1985, B. Aukema.
76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
450. Tropidophlebia costalis
References. - Reclaire (1932: 100, as Camptotelus li-
neolatus Schilling; 1943: 109, as C. costalis).
Published records. - ® Utrecht: Soest (Reclaire 1943).
® N-Holland: Hilversum (Reclaire 1932).
Present status. - Not recorded since 1943: Soest, 9.vi,
A. Reclaire (ZMA).
454. Plinthisus pusillus (fig. 99)
References. - Reclaire (1932: 105, 258; 1948: 42; 1951:
a).
Published records. - © Drenthe: Roden (Reclaire
1948). ® Gelderland: Otterlo (Reclaire 1932). ® N-Hol-
land: Aerdenhout, Hilversum (Reclaire 1932); Texel, De
Koog (Reclaire 1951). ® Z-Holland: Wassenaar, Katwijk
(Reclaire 1932). ® N-Brabant: Breda (Reclaire 1932).
Unpublished records. - ® N-Holland: Hilversum,
22.ix.1934, 26.v.1935, 3.ix.1938 & 20.viii.1940, A. Re-
claire (ZMA); Laren, 3.iii.1935, A. Reclaire (ZMA). € Z-
Holland: Oostvoorne, 24.vii.1956, R. H. Cobben; Noord-
wijk, 13.ix.1987, 1 4, K. den Bieman. ® Gelderland:
Hoenderloo, 5.ix.1982, 1 ®, K. den Bieman.
455. Tropistethus holosericeus (fig. 100)
References. — Reclaire (1932: 103; 1934: 50; 1936: 247;
1940: 107); Blöte (1954: 84).
Published records. - © Utrecht: Rhenen (Reclaire
1940). ® Z-Holland: Wassenaar (Reclaire 1936). ® Lim-
burg: Maastricht (Reclaire 1932 1934; Blöte 1954); Be-
melen (Reclaire 1936); Eijs (Reclaire 1940).
Unpublished records. - © Limburg: Bemelen,
10.ix.1966, P. J. Brakman (P); Colmont, 14.v.1966 &
1.vii.1967, Mheer, 18.vii.1968 & 9.v.1970, P. Poot.
456. Drymus latus
References. — Reclaire (1932: 115, as D. confusus;
1936: 248).
Published records. - ® Limburg: Maastricht (Reclaire
1932); Bemelen, Eijs, Schin op Geul, Wijlre (Reclaire
1936).
Unpublished records. - ® Limburg: Colmont,
7.iv.1963, 1 6, P. J. Brakman (RML).
457. Drymus pilicornis
References. — Fokker (1884); Reclaire (1951: 8).
Published records. - ® N-Holland: Haarlem, ix, Groll
(RML) (Fokker 1884); Heemstede, 31.vii.1942, 1 @, D.
L. Uyttenboogaart (ZMA) (Reclaire 1950).
Present status. - Not recorded since 1942.
459. Drymus pumilio
References. - Aukema (1986a).
Published records. - € Zeeland: Cadzand (Aukema
1986a). ® Limburg: Maastricht, Mechelen (Aukema
1986a).
Present status. - Local, very rare. Three 9 were collec-
ted at different localities between 1921 and 1960.
Fig. 98. Oxycarenus modestus.
Fig. 99. Plinthisus pusillus.
AUKEMA:
Fig. 100. Tropistethus holosericeus.
Fig. 101. Drymus reyis.
Checklist Hemiptera-Heteroptera Vi
460. Drymus ryeii (fig. 101)
References. - Cobben (1958a: 5, as D. picinus Rey).
Present status. - Widespread and common.
464. Eremocoris podagricus
References. — Reclaire (1932: 117; 1936: 248; 1940:
108; 1948: 42; 1951: 8); Brakman (1952: 130); Blöte
(1954).
Published records. - ® Limburg: Maastricht (Reclaire
1932, 1948; Blöte 1954); Valkenburg, Wijlre (Reclaire
1936); Bunde (Reclaire 1940); Bemelen (1951). ® Zee-
land: Oostkapelle (Brakman 1952).
Unpublished records. - © Limburg: Vaals, 10.v.1964 &
Bemelen, 10.ix.1966, P. Poot; Oud-Valkenburg, Gerendal,
6.vi, l.vii & 16.7.1977, C. Dolleman.
468. Lamproplax picea
References. — Reclaire (1943: 110, as Drymus piceus;
1948: 42); Aukema & Alderweireldt (1989, in press).
Published records. — ® Friesland: Schiermonnikoog
(Aukema & Alderweireldt 1989). ® Drenthe: Wijster
(Aukema & Alderweireldt 1989). ® Gelderland: Winters-
wijk (Reclaire 1943). ® Limburg: Haelen (Reclaire
1948); Herkenbosch, Meijnweg (Aukema & Alderwei-
reldt 1989)
472. Scolopostethus pilosus
References. - Reclaire (1932: 119; 1943: 110).
Published records. - © Gelderland: Varsseveld (Re-
claire 1932); Winterswijk (Reclaire 1943).
Unpublished records. — © Overijssel: Kalenberg,
Weerribben, 20.v.1978, 1 ®, C. Dolleman.
473. Scolopostethus pseudograndis f.
nov. spec.
Note. - Miss B. Berkhout discovered this species revi-
sing the Scolopostethus material in the Leiden Museum
(RML).
Records. — © Limburg: Maastricht, St. Pietersberg,
16.viii.1950, 1 6, 17.x.1950, 2 4, 1 Q & Gronsveld, Rie-
senberg, 20.x.1950, 1 9, H. C. Blöte et al. (RML).
Present status. - Not recorded since 1950.
477. Taphropeltus hamulatus
References. - Reclaire (1932: 119; 1936: 249; 1940:
108).
Note. - the specimen from Hilversum (Reclaire 1940)
belongs to T. contractus.
Published records. - ® Zeeland: Zierikzee (Reclaire
1932). © Limburg: Eijs (Reclaire 1932); Schin op Geul
(Reclaire 1936).
Unpublished records. - ® Limburg: Wijlre, 11-
12.viii.1958 & 10.viii.1959, W. H. Gravestein (ZMA);
Colmont, á.iv.1959, W. Hellinga (ZMA); Ibid., 6.iv.1965,
P. Poot (ZMA) & 8-9.iv & l.vii.1967, P. Poot; Mheer,
9.v.1970, P. Poot.
479. Lasiosomus enervis
References. — Reclaire (1940: 107); Brakman (1952:
130).
78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Published records. — ® Zeeland: Oostkapelle,
27.vi.1949, P. J. Brakman (RML) (Brakman 1952). ®
Limburg: Eijs, 13.ix.1936, 1 8, A. Reclaire (ZMA) (Re-
claire 1940).
Unpublished records. - © Limburg: Vaals, 13.vi. 1956,
1g, 1 9, W. H. Gravestein (ZMA).
Present status. - Not recorded since 1956.
484. Pachybrachius luridus (fig. 102)
References. — Fokker (1886: 298, as Plociomerus luri-
dus; Reclaire (1932: 101; 1936: 247; 1943: 109).
Published records. - ® Drenthe: Gasteren (Reclaire
1936); Wijster (Reclaire 1943). ® Gelderland: Winters-
wijk (Fokker 1886); Arnhem (Reclaire 1932). ® Utrecht:
Baarn, Vinkeveen (Reclaire 1932). ® N-Holland: Anke-
veen (Reclaire 1932); Kortenhoef (Reclaire 1936).
Unpublished records. — © Gelderland: Tongeren,
22.1x.1983, 1 À, 1 9, K. den Bieman (AU, BI). ® N-
Holland: Kortenhoef, 16.vi.1951, H. Smit (ZMA).
485. Aellopus atratus
References. - Reclaire (1932: 109); Cobben (1958: 16).
Published records. - ® Limburg: Bemelen (Reclaire
1932); Schin op Geul (Reclaire 1936; Cobben 1958).
Unpublished records. — © Limburg: Bemelen,
23.ix.1949, R. Batten (P); Ibid., 15.viti.1966, P. Poot &
10.ix.1966, P. Poot (ZMA); Valkenburg, 22.vi.1965, P.
Poot.
488. Peritrechus angusticollis
References. — Reclaire (1936: 248).
Published records. — ® Limburg: Schinveld,
24.x11.1934, 1 9, J. Cremers (NMM) (Reclaire 1936).
Unpublished records. - ® Limburg: Belfeld, 22.vi.1985,
1 6, H. Peeters (P).
490. Peritrechus lundii (fig. 103)
References. — Reclaire (1932: 108, as P. sylvestris;
1936: 248; 1940: 107; 1948: 42; 1951: 8).
Present status. - Formerly widespread, but rare. Col-
lected for the last time in 1951: Vogelenzang, Amster-
damse Waterleidingduinen, 19.v (ZMA).
495. Raglius alboacuminatus
References. - Cobben (1948: 50; 1953: 186-187).
Published records. - ® Limburg: Vlodrop, Roermond
(Cobben 1948, 1953).
Present status. - Not recorded since 1951: Vlodrop, St.
Ludwig, 17.viii, R. H. Cobben (AU, CB, LUW).
496. Megalonotus antennatus
References. - Fokker (1886: 298, as Rhyparochromus
antennatus), Reclaire (1932: 101, 258, as Rhyparochro-
mus), Aukema (1983).
Note. - The specimen mentioned by Fokker (1886)
from Vorden (ix, Groll), most likely concerns the speci-
men from Warnsveld (3.ix.1885, leg. Groll) in the Am-
sterdam Museum.
Published records. — © Gelderland: Warnsveld (Re-
claire 1932). © Limburg: Schin op Geul (Reclaire 1932);
Wahlwiller (Aukema 1983).
Fig. 102. Pachybrachius luridus.
Fig. 103. Peritrechus lundi.
AUKEMA: Checklist Hemiptera-Heteroptera 79
Unpublished records. - © Friesland: Schiermonni-
koog, 14.v.1961, G. Stobbe (FNM). ® Limburg: Colmont,
3.viii.1960, 11.v.1963 & 11.v.1965, Br. Arnoud (CB); Ibid.,
6.iv & 3.vii.1965, 15.x.1968, P. Poot (P, ZMA); Ibid,
ix.1969, C. Berger (P); Ibid, Wrakelberg, 6.v.1977 &
7.v.1979, C. Dolleman; Wahlwiller, Kruisberg, 16 &
28.vii.1977, C. Dolleman.
500. Megalonotus sabulicola (fig. 104)
References. — Fokker (1884: 117); Reclaire (1932: 103;
1934: 50; 1936: 247; 1951: 7).
Present status. - Widespread, but restricted to sandy
soils.
503. Emblethis griseus
References. - Snellen van Vollenhoven (1878a: 96-97,
as Pachymerus griseus); Brakman (1952: 130, as E. ver-
basci).
Published records. - ® Zeeland: Walcheren (Snellen
van Vollenhoven 1878; Fokker 1884, as verbasci; Reclaire
1932, as verbasci); Domburg, 6.vii-18.viii.1951, P. J. Brak-
man (RML, P) (Brakman 1952).
Unpublished records. - ® Zeeland: Domburg,
1.vii.1983, 12 & 14.vii.1986, B. Aukema.
Present status. - Uncertain, because in 1987 the locality
was destroyed when the dykes were raised.
504. Emblethis verbasci
Fig. 104. Megalonotus sabulicola. References. - Reclaire (1932: 114).
Published records. - © N-Brabant: Bergen op Zoom
(Reclaire 1932). ® Limburg: Schin op Geul (Reclaire
1932).
m Present status. - Not recorded since 1944: Schin op
Geul, vii, 1 6, A. Reclaire (ZMA).
510. Trapezonotus desertus (fig. 105)
References. - Woodroffe (1960).
Present status. - Common on heather heaths.
513. Dicranocephalus agilis
References. - Cobben (1948: 50); Blöte (1954: 84).
Note. - The specimens recorded from Arnhem, cited
by Fokker (1883) belong to D. medius and the one from
Den Haag (Snellen van Vollenhoven 1878) is not trace-
able (Cobben 1948).
Published records. — © Limburg: Vlodrop (Cobben
1948); Maastricht (Blöte 1954).
Unpublished records. - © N-Holland: Zaandam,
25.vii.1963, 1 9, ex larva, J. H. Woudstra (specimen of
doubtful origin). ® N-Brabant: Woensdrecht, 26.ix.1955,
1 9, P.J. Brakman.
514. Dicranocephalus medius
References. - Fokker (1883: 246-247, as D. agilis);
Cobben (1948: 50); Gravestein (1952a: cxvi); Blöte
(1954: 84); Loof (1956).
Note. - One unlabelled though probably Dutch speci-
men in the collection of the Dutch Entomological Society
(Fokker 1883).
Published records. - ® N-Holland: Vogelenzang (Loof
Fig. 105. Trapezonotus desertus. 1956). ® Gelderland: Arnhem (Fokker 1883, as agilis;
80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Fig. 106. Gonocerus acuteangulatus. Fig. 107. Gonocerus juniperi.
Fig. 108. Bathysolen nubilus. Fig. 109. Corizus hyoscyami.
AUKEMA:
Fig. 110. Rhopalus subrufus.
Fig. 111. Stictopleurus abutilon abutilon.
Checklist Hemiptera-Heteroptera 81
Cobben 1948). ® Limburg: Vlodrop (Cobben 1948), Rij-
kel (Gravestein 1952a), Maastricht (Blöte 1954).
Unpublished records: ® Limburg: Rijkel, 17.vi.1953, P.
J. Brakman (P); Swalmen, 20.vi.1965, 1 4, P. J. Brakman
(P).
515. Gonocerus acuteangulatus (fig. 106)
References. — Snellen van Vollenhoven (1878a: 60-61,
as G. venator); Reclaire (1932: 256; 1951: 4, as G. juniperi
from Herkenbosch); Cobben (1953: 174; 1958: 15-16).
Published records. — ® Gelderland: Arnhem, Ooster-
beek (Snellen van Vollenhoven 1878). ® Limburg: Hou-
them, Bunde, Venlo (Snellen van Vollenhoven 1878);
Herkenbosch, Vlodrop, Boukoul (Cobben 1953); Bunde
(Reclaire 1932).
Present status. - Not recorded since 1950: Vlodrop,
27.vii, R. H. Cobben.
516. Gonocerus juniperi (fig. 107)
References. - Reclaire (1932: 84; 1951: 4); Gravestein
(1943: viii).
Note. - The record from Herkenbosch (Reclaire 1950)
concerns G. acuteangulatus (Cobben 1958).
Published records. - ® Overijssel: Ommen (Reclaire
1932); Lemelerberg (Gravestein 1943). ® N-Brabant:
Lierop, Deurne (Reclaire 1951).
Unpublished records. - © Limburg: Venray, 28.viii-
25.1x.1951, R. H. Cobben (CB, LUW, NMM).
Present status. - Not recorded since 1951.
522. Arenocoris waltlii
References. - Snellen van Vollenhoven (1878a: 66-67,
as Pseudophloeus waltlii), Fokker (1891: 357, as Pseudo-
phlaeus waltlii), Reclaire (1932: 86, 256, as Pseudo-
phloeus waltlt).
Published records. - ® Gelderland: Nijmegen (Fokker
1891); Buren (Reclaire 1932). ® Zeeland: Walcheren
(Snellen van Vollenhoven 1878); Vlissingen (Reclaire
1932).
Present status. - Not recorded since 1892: Buren, vii,
H. J. Veth (RML).
523. Bathysolen nubilus (fig. 108)
References. — Reclaire (1936: 246; 1948: 40); Blöte
(1954: 84).
Published records. - © Limburg: Bemelen, Schin op
Geul (Reclaire 1936); Roermond (Reclaire 1948); Maas-
tricht (Blöte 1954).
Unpublished records. - © Limburg: Kerkrade, 14. viii
& 1.1x.1936, Willemse (ZMA); Schinveld, 30.viii. 1936,
Willemse (ZMA); Schin op Geul, 10.viii.1941, M. van den
Boorn (P); Vlodrop, 23.v & 20.vii.1948; Heel, 14.ix.1953,
1 g, R. H. Cobben; Venlo, 15.ix.1982, B. van Aartsen
(ZMA); Ibid., 17.vi.1982 & 21.v.1983, B. Aukema; Ibid.,
3.viii.1983, B. Aukema & J. H. Woudstra; Maastricht, St.
Pietersberg, 18.viii & 2.ix.1983, 29.viii.1984, B. Aukema.
527. Liorhyssus hyalinus
References. - Reclaire (1936: 246).
Published records. — © Gelderland: Nunspeet,
viii.1923, 1 9, D. MacGillavry (ZMA) (Reclaire 1936).
82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Present status. - Not recorded since 1923.
528. Corizus hyoscyami (fig. 109)
References. - Reclaire (1932: 89, as Therapha hyosci-
ami; 136: 246; 1948: 41; 1951: 5).
Present status. - Widespread, but rare. Not recorded
between 1949 and 1977. Since then only in Limburg:
Gronsveld (J. H. Woudstra); Kunrade, Maastricht, Vlo-
drop (B. van Aartsen, ZMA).
530. Rhopalus subrufus (fig. 110)
References. - Reclaire (1932: 90, 257, as Corizus subru-
fus), 1934: 49; 1940: 106; 1943: 108; 1951: 5).
Published records. — @ Friesland: locality unknown
(Reclaire 1932). © Utrecht: Driebergen (Reclaire 1932).
® N-Holland: Velsen (Reclaire 1951). ® Limburg:
Eijsden, Schin op Geul (Reclaire 1932); Houthem (Re-
claire 1932); Gronsveld (Reclaire 1934); Wijlre (Reclaire
1940); Valkenburg (Reclaire 1932, 1943); Geulhem (Re-
claire 1951).
Unpublished records. - @ N-Holland: Heemskerk,
14.x.1951, J. H. Woudstra; Ibid., 16.v.1959, J. Beernink. ©
Limburg: Maastricht, St. Pietersberg, 12.vi.1935 (NMM);
Ibid, 18.vii.1983 & 29.viii.1984, B. Aukema; Roermond,
15.v.1949, 1 9, R. H. Cobben; Vlodrop, 22.ix.1951, R. H.
Cobben.
534. Stictopleurus abutilon abutilon (fig. 111)
References. — Cobben (1948: 50; 1951: 53); Reclaire
(1951: 5); Brakman (1952: 130); Blöte (1954: 84).
Published records. - ® Gelderland: Kootwijk (Reclaire
1951). ® Zeeland: Oostkapelle (Brakman 1952). ® N-
Brabant: Eindhoven (Reclaire 1951). © Limburg: Beesel,
Boukoul, Linne, Melick, Herkenbosch, Roermond, St.
Odiliënberg, Vlodrop (Cobben 1948; Reclaire TOSI
Haelen (Cobben 1951); Maastricht (Blöte 1954).
Unpublished records. - ® Gelderland: Nijmegen,
31.viti.1986, D. J. Hermes; Pannerden, 19.vi.1977, C. Dol-
leman. ® Utrecht: Maartensdijk, 28.vii.1950, W. H. Gra-
vestein (ZMA). ® N-Brabant: Best, 7.ix.1950, M. van den
Boorn (P); Ibid., 9.x.1976 & 24.viii.1974, H. W. van der
Wolf (AU); Eindhoven, Ekkersrijt, 8.x.1976, H. W. van
der Wolf (AU). ® Limburg: Heerlen, 25.vi.1949, Br.
Arnoldus (ZMA); Echt, 16.viii.1949, R. H. Cobben; Me-
lick-Herkenbosch, 26.viii & 9.ix.1949, 21.ix.1951 & 14.ix.
1953, R. H. Cobben; Koningsbosch, 21.v.1949, R. H.
Cobben; Maastricht, St. Pietersberg, 26.ix.1983, 2.ix &
15.x.1984, B. van Aartsen (ZMA); Ibid, 8-11.vii &
2.1x.1983, B. Aukema; Mook, 21.viii.1983, K. den Bieman;
Schaesberg, 1.v & 22.vi.1987, G. R. langohr (RML).
Present status. - Widespread, but rare. Collected in two
distinct periods: 1947-1953 and since 1976.
535. Stictopleurus punctatonervosus
(fig. 112)
References. — Blöte (1950: 19-20; 1954: 84); Cobben
(1951: 53); Arnoud (1958).
Published records. — ® Limburg: Maastricht (Blöte
1950, 1954); Melick-Herkenbosch (Cobben 195 1), Bruns-
sum (Arnoud 1958); Bemelen, Cadier, Keer (Blöte 1954).
Unpublished records. — @ Gelderland: Nijmegen,
Fig. 112. Srictopleurus punctatonervosus.
Fig. 113. Aethus nigrita.
AUKEMA:
459. Drymus pumilio. Scale: 1 mm.
31.viii.1986, 17.iv & 19.vii.1987, D. J. Hermes; Ibid.,
6.xi.1987, B. Aukema; Tolkamer, 9.viii.1988, G. Morseld.
© N- Brabant: Best, 4.vi.1983, B. Aukema; Oirschot,
12.x.1985, B. van Aartsen (ZMA). ® Limburg: Vlodrop,
22.1x.1951 & 2.viii.1954, R. H. Cobben; Ibid., 10.vii.1981
& 1.1x.1984, B. van Aartsen (ZMA); Maastricht,
l.viii.1972, 26-27.ix.1983, 17 & 21.vii, 18.viii, 2.ix &
15.x.1984, B. van Aartsen (ZMA); Ibid., 22.ix.1975,
24.viii & 14.ix.1983, 29.viii.1984 and 7.vi.1985, B. Au-
kema; Venlo, 17.vii.1979, 17.vi.1980, 7.vii & 15.ix.1982,
22.vi, 16.vii & 1.x1.1984, B. van Aartsen (ZMA); Ibid.
30.x.1982, B. Aukema & 3.viii.1983, B. Aukema & J. H.
Woudstra; Colmont, 27.ix.1983, B. van Aartsen (ZMA);
Nijswiller, 19.viii.1987, G. R. langohr (RML).
541. Sehirus morio
References. - Fokker (1883: 241); Reclaire (1932: 65);
Cobben (1953: 172).
Published records. - ® Utrecht: Bilthoven (Reclaire
1932). ® Gelderland: Arnhem (Fokker 1883; Reclaire
1932). ® Limburg: Montfort (Cobben 1953).
Unpublished records. - © Utrecht: Maartensdijk,
Groenekan, 2.vii.1950, W. J. Kabos (ZMA).
Present status. - Not recorded since 1950.
543. Aethus nigrita (fig. 113)
References. — Reclaire (1932: 63, as Cydnus nigrita;
1936: 244; 1951: 2); Cobben (1953: 172); Brakman (1960:
24).
Checklist Hemiptera-Heteroptera 83
Published records. - ® Gelderland: Nijmegen, Putten,
Wageningen (Reclaire 1932); Laag-Soeren (Reclaire
1936). ® N-Holland: Hilversum (Reclaire 1932, 1936).
®© Z-Holland: ‘s-Gravenzande (Reclaire 1951). © Zee-
land: Zierikzee (Reclaire 1932); Zoutelande (Brakman
1960). © N-Brabant: Uden (Reclaire 1932). © Limburg:
Maasniel, Melick, Roermond (Reclaire 1951); Rijkel
(Cobben 1953).
Unpublished records. — @ Gelderland: Arnhem,
19.v.1898 (ZMA); Ibid., 20.iv, G. A. Six (RML); Velp,
2l.iv & 2-4.v.1901 (ZMA); Wageningen, 19.vi.1951
(LUW). e N-Brabant: Bergen op Zoom, 2.ix.1962, P. J.
Brakman (P). ® Limburg: Heijthuizen, 16.iv.1950, C.
Berger (P); Lomm, 25.viii.1968, P. Poot; Venlo, 23-
25.v1.1981, B. Aukema.
544. Cydnus aterrimus
References. — Gravestein (1952a: cxviii); Cobben
(1953: 177).
Published records. - ® Limburg: Rijkel, 1.vii.1951, W.
H. Gravestein, R. H. Cobben & P. J. Brakman (Gravestein
1952a).
Present status. - Not recorded since 1951.
547. Odontoscelis lineola
Synonym: O. dorsalis auct. nec Fabricius.
References. - Göllner-Scheiding (1986) showed that
the West European specimens recorded as Odontoscelis
dorsalis in fact belong to O. lineola Rambur, whereas the
true dorsalis is a southern species, not occurring on the
European continent.
548. Phimodera humeralis
References. - Snellen van Vollenhoven (1878a: 19-20,
as P. galgulina), Reclaire (1932: 68; 1936: 245; 1943:
107).
Published records. - ® Utrecht: Soest (Reclaire 1943).
® N-Holland: Bergen aan Zee (Reclaire 1936). © Z-
Holland: Scheveningen (Snellen van Vollenhoven 1878;
Reclaire 1932).
Unpublished records. - © Gelderland: Apeldoorn,
1901, 1 9 (ZMA).
Present status. - Not recorded since 1941: Soest, 4.v,
R. Tolman (ZMA).
549. Eurygaster austriaca (fig. 114)
References. - Snellen van Vollenhoven (1878a: 15, as
Tetyra hottentotta F.), Fokker (1883: 240, as E. hotten-
totta); Reclaire (1932: 69; 1936: 245).
Published records. - ® Gelderland: Velp (Fokker
1883). © Utrecht: Utrecht (Fokker 1883). © N-Holland:
Overveen (Snellen van Vollenhoven 1878); Haarlem
(Fokker 1883). ® Z-Holland: Den Haag (Snellen van
Vollenhoven 1878). ® Zeeland: Walcheren, Zierikzee
(Fokker 1883); Westenschouwen (Reclaire 1936). ®
Limburg: Venlo (Reclaire 1932); Schin op Geul (Reclaire
1936).
Present status. - Not recorded since 1935: Schin op
Geul, 15.ix, A. Reclaire (ZMA).
84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
552. Graphosoma lineatum (fig. 115)
References. — Snellen van Vollenhoven (1878a: 17, as
Trigonosoma nigrolineata Rossi); Blöte (1950a); Cobben
(1958: 15); Arnoud (1958).
Published records. - © N-Brabant: without locality
(Snellen van Vollenhoven 1878). ® Limburg: Amby, Be-
melerberg, 1948, D. Piet; Maastricht, Cannerbos,
19.v.1950, 1 9, C. v. Heijningen (RML) (Blöte 1950a);
Echt,17.vi.1951, 1 g, 1 9, R. Geurts (CB), Epen, begin
vi.1953, V. Westhoff et al. (Cobben 1958); Terwinselen,
15.v.1948 & Heerlen, 28.vu.195 1, Br. Arnoud & J. van Loo
(Arnoud 1958).
Unpublished records. — © Limburg: Bemelen,
19.viii. 1965, Br. Virgilius (P).
560. Rubiconia intermedia
References. - Reclaire (1932: 73; 196: 245); Cobben
(1958a: 3-4).
Published records. - € Gelderland: Vierhouten (Re-
claire 1932); ? Apeldoorn (Reclaire 1936); Ede (Cobben
1958a).
Unpublished records. — © Overijssel: Nijverdal,
5.x.1972, B. van Aartsen (ZMA). ® Gelderland: Gortel,
4-6.x.1983, 16 & 23.x.1983, B. van Aartsen (ZMA, P);
Ibid., 6.x & 11.xi.1984, B. van Aartsen (P); Hoog Soeren,
2.1x.1984, B. van Aartsen (ZMA, P); Ibid., 15.xi. 1984, B.
van Aartsen (P); Terlet, 29.vii, 15.viii & 28.ix.1984, B. van
Aartsen (P).
561. Holcostethus sphacelatus
References. - Reclaire (1932: 74, as Peribalus sphace-
latus).
Published records. — © Limburg: Zuid-Limburg,
vi.1909, 1 4, D. L. Uyttenboogaart (ZMA) (Reclaire
1932).
Unpublished records. - ® Limburg: Maastricht, St.
Pietersberg, 26.ix.1983, 1 specimen, B. van Aartsen
(ZMA). Furthermore one 9 labelled ‘Holland, v. Voll.’
(RML).
562. Holcostethus vernalis (fig. 116)
References. — Reclaire (1932: 74, as Peribalus vernalis;
1936: 245; 1940: 105; 1943: 107; 1951: 3); Brakman
(1952: 131); Cobben (1953: 173).
Published records. - ® Z-Holland: Meijendel (Reclaire
1932). ® Zeeland: Oostkapelle (Brakman 1952). ® Lim-
burg: Zuid-Limburg (Reclaire 1932); Heek (Reclaire
1936); Schin op Geul (Reclaire 1940); Bemelen (Reclaire
1943); Vlodrop (Reclaire 1951; Cobben 1953).
Unpublished records. - ® N-Holland: Vogelenzang,
2.vi.1957, J. van der Wiel (ZMA). © Limburg: Bunde,
ix.1936 (NMM); Vlodrop, 30.vi.1951, W. H. Gravestein
(ZMA); Ibid., P. J. Brakman (P); Maastricht, St. Pieters-
berg, 1.vi.1951, A. A. W. Smit (ZMA); Ibid., 2.ix.1983, B.
Aukema; Ibid., 26.ix.1983, 2.ix & 15.x.1984, 30.v, 11 &
31.1x.1985, B. van Aartsen; P. Vijlen, 23.v.1953, P. van der
Wiel (ZMA); Terziet, 19.v.1966, P. Poot; Schaesberg,
22.vi.1987, G. R. Langohr (RML).
Fig. 114. Ewrygaster austriaca.
Fig. 115. Graphosoma lineatum.
AUKEMA:
Checklist Hemiptera-Heteroptera 85
PF?
Fig. 116. Holcostethus vernalis.
Fig. 118. Chlorochroa pinicola.
Fig. 117. Palomena viridissima.
564. Palomena viridissima (fig. 117)
References. — Reclaire (1932: 74, as ‘widespread
throughout the country’).
Present status. - Formerly widespread, but not recor-
ded since 1952: Arnhem, 9.v. (RINA).
565-566. Chlorochroa
Synonym: Pitedia Reuter. (see Vinukorov et al. 1988).
566. Chlorochroa pinicola (fig. 118)
References. - Reclaire (1932: 75; 1940: 105; 1948: 39-
40); Gravestein (1952a: cxv); Leston (1952: 84); Brak-
man (1960: 24); Kleinhout (1962: 39).
Published records. - ® Gelderland: Oosterbeek, Arn-
hem (Reclaire 1932); Leuvenum (Reclaire 1940); Hoge
Veluwe (Gravestein 1952; Leston, 1952; Kleinhout
1962). ® Utrecht: Doorn, Maarn (Reclaire 1948). © N-
Holland: Hilversum (Reclaire 1948). © Limburg: Venlo
(Reclaire 1932); Melick-Herkenbosch, Beegden, Heel
(Reclaire 1948). e Zeeland: Vrouwenpolder (Brakman
1960).
Unpublished records. - ® Gelderland: Hoge Veluwe,
28.11.1952, 1 @ (RINA). ® Utrecht: Groenekan,
11.1x.1944, R. Buisman (ZMA); Hollandsche Rading,
14.v.1952 & iv.1953, W. H. Gravestein (ZMA). ® N-
Brabant: Breda, 7.viii.1950, M. van den Boorn (P); Gel-
drop, 6.ix.1983, 1 9, J. de Jonge (PDW); Baronie Cra-
nendonck, 12.vi.1987, 1 9, G. R. Langohr (RML). ®
Limburg: Haelen, viii,1950, C. Willemse (ZMA).
Present status. - Recorded only twice since 1953.
86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
567. Carpocoris fuscispinus
References. - Fokker (1883: 242, as C. nigricornis);
Reclaire (1932: 75-76; 1940: 105; 1943: 107).
Published records. - € Gelderland: Beekhuizen (Fok-
ker 1883); Oosterbeek (Reclaire 1932). ® N-Brabant:
Burgst (Reclaire 1932). © Limburg: Schin op Geul (Re-
claire 1940); Arcen (Reclaire 1943).
Unpublished records. - ® N-Brabant: Best, viii. 1950,
M. van den Boorn (P).
Present status. - Not recorded since 1950.
572. Eurydema dominulus (fig. 119)
References. - Reclaire (1932: 77; 1936: 246; 1940: 105;
1948: 40); Cobben (1953: 173).
Published records. — ® Overijssel: Denekamp, Oot-
marsum (Reclaire 1932); Tubbergen (Reclaire 1948). ®
N-Brabant: Breda (Reclaire 1932). ® Limburg: Meers-
sen, Schinnen, Spaubeek, Venlo, Valkenburg (Reclaire
1932); Maastricht (Reclaire 1936); Houthem (Reclaire
1936, 1940). Vlodrop (Cobben 1953).
Unpublished records. - © Overijssel: Mander,
20.vii.1944, D. Piet et al. (P); Vasse, 1.viii.1947, D. Piet
(ZMA); Ootmarsum, vii.1947, 1 9, G. Stobbe (FNM),
Ommen, vi.1964, W. H. Gravestein (ZMA); Ibid.
ix.1967, W. Hellinga (ZMA). ® N-Brabant: Best,
1.vi.1983, 1 4, B. van Aartsen (ZMA).
574. Eurydema ventralis
References. — Reclaire (1932: 76, as E. ornatum (Lin-
naeus)).
Published records. — ® N-Brabant: Surroundings of
Breda, 1892, 1 & & 1893, 1 9, Smits van Burgst (ZMA)
(Reclaire 1932).
Present status. - Not recorded since 1893.
579. Jalla dumosa (fig. 120)
References. - Snellen van Vollenhoven (1878a: 25-26,
as Asopus dumosus), Fokker (1883: 244-245); Reclaire
(1943: 108; 1951: 4); Brakman (1960: 24).
Published records. - © Gelderland: Beekhuizen (Re-
claire 1943). ® Utrecht: Baarn (Reclaire 1951). ® N-
Holland: Driebergen (Snellen van Vollenhoven 1878). ®
Zeeland: Walcheren (Snellen van Vollenhoven 1878);
Zierikzee (Fokker 1883); Hoek (Brakman 1960).
Unpublished records. — © Friesland: Terschelling,
2.viii.1953 & 20r-23.vii. 1954, W. H. Gravestein (ZMA);
Ibid, 17.viii.1953, P. E van Heerdt; Ibid, Lies,
20.viii.1957, W. H. Gravestein (ZMA); Ibid., Oosterend,
5-15.viii.1969, F Meerman. @ Limburg: Montfort,
14.x.1951, 1 Q (CB).
Present status. - Recorded only once since 1957.
CONCLUDING REMARKS
Despite the relatively low collecting efforts
through the years (see for instance figs. 72 and 73,
the distribution maps of Trigonotylus caelestiahum
and T. ruficornis, two of the very common species)
the total number of species (586) recorded from
The Netherlands is high in comparison with data
from other West European countries: Sweden
-
Fig. 119. Eurydema dominulus.
Fig. 120. Jalla dumosa.
AUKEMA: Checklist Hemiptera-Heteroptera 87
Table 1. Total number of species per family and per region
FAMILY REGION
ERIN GARY IDI MON ET MN GIL AUTONET ZZE TOMS ZZE N 03) INIL
1 Dipsocoridae 1 0 i 0 l l 1 1 1 0 0 2 2
2 Nepidae 2 2 2 2 2 2 2 2 2 2 2 2 2
3. Corixidae 25 108237574 IOM E25 TE a Te EE OI 22 5
4 Naucoridae l 1 1 1 1 I 2 1 2 1 2 1 2
5 Aphelocheiridae 0 0 0 1 0 0 0 0 0 0 1 1 1
6 Notonectidae 6 4 6 6 3 6 5 5 4 3 5 5 6
7, Pleidae Il 1 1 1 1 Il Il 1 1 1 1 1 1
8 Mesoveliidae l 0 1 1 0 1 1 1 1 1 I 1 l
9 Hebridae 2 2 2 2 2 2 2 2 2 2 2 2 2
10 Hydrometridae 1 2 2 2 1 2 2 2 2 I 2 2 2
11 Veliidae 2 1 2 3 2 4 3 3 3 2 4 5 5
12 Gerridae 5 5 7 8 4 9 U 8 7 6 9 9 9
13 Saldidae 15 8 9 12 8 12 10 14 10 10 13 Bs). 20)
14 Tingidae 10 3 12 7 3 23 TO RZ: 13 10 IC 23 30
15. Microphysidae 5 1 5 5 0 6 5 5 5 3 3 5 6
16 Miridae 1242 1 53) 136) A479) vO AIG) ASG 7 A 2 2 Ene 200) 05231
17 Nabidae 10 6 10 10 5 12 il 11 10 13 12 13 13
18 Anthocoridae 15 7 20 16 11 27 DO 17 17 17 30 32
19 Cimicidae 1 1 0 1 0 3 3 3 3 3 2 2 5
20 Reduviidae 3 1 6 4 1 8 5 5 4 4 6 6 9
21 Aradidae 3 0 4 2 1 5 3 3 3 2 4 4 6
22 Piesmatidae 1 0 2 1 0 1 1 3 2 2 2 2 3
23 Berytidae 4 0 3 2 0 6 4 5 5 7 5 8 9
24 Lygaeidae 44 NOA SS Ae GI a O > Aye OS
25 Pyrrhocoridae 0 0 0 0 0 1 1 1 l 1 il 1 1
26 Stenocephalidae 0 0 0 0 0 1 0 1 0 0 1 2 2
27 Coreidae 4 1 2 3 1 df 5 6 6 Ÿ 7 9 11
28 Alydidae 0 0 0 0 0 1 1 1 0 0 1 1 1
29 Rhopalidae 4 1 4 4 2 9) 7 4 3 5 7 8 9
30 Cydnidae 4 1 3 2 0 7 6 Vi 6 5 5 8 9
31 Thyreocoridae l 0 0 0 0 1 1 l 1 0 1 1 1
32 Scutelleridae 1 0 0 0 0 6 5 6 5 4 3 4 6
33 Pentatomidae 14 5 15 16 AE 25 Allee ZO 11 23 vol 200029
34 Acanthosomatidae 5 3 5 5 3 6 6 5 3 3 6 6 6
Number of species 315 144 324 326 160 476 399 451 337 321) 391° 496) 586
(577), Norway (395), Denmark (492), North West
Germany (585) and the British Isles (532) (Ander-
sen & Gaun 1974; Coulianos & Ossiannilsson 1976;
Southwood & Leston 1964; Wagner & Weber
1967).
From table 1 it is also clear that the collecting
effort has been different for the various regions.
Limburg, Gelderland and Noord-Holland for ins-
tance are far ahead concerning number of species
collected, whereas Groningen lays back considera-
bly. The number of species collected in the recently
reclamed polders of Flevoland (Oostelijk and
Zuidelijk Flevoland ran dry in 1957 and 1968 re-
spectively) is still relatively low (151), but rapidly
increasing. The number of species collected in
Friesland (314) is largely due to the high collection
efforts on the West Frisian Islands of Vlieland,
Terschelling, Ameland, Terschelling, and Schier-
monnikoog, where 258 species were collected.
From the Frisian mainland thusfar only 197 species
were recorded.
The list contains 48 species which were not re-
corded since 1960. Of these Galeatus maculatus,
Acetropis gimmerthalii, Phymata crassipes, Spilo-
stethus saxatilis, Geocoris ater, Arenocoris waltli
and Eurydema ventralis were collected only before
1900. The following 18 species were collected since
1960 only: Sigara iactans, S. longipalis, Deraeocoris
flavilinea, Tytthus geminus, Brachyarthrum limita-
tum, Psallus assimilis, P. pseudoplatani, Criocoris
88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
sulcicornis, Reuteria marqueti, Orthotylus fusces-
cens, Capsodes sulcatus, Elatophilus nigricornis,
Anthocoris amplicollis, Scoloposcelis pulchella,
Aradus signaticornis, Orsillus depressus, Kleidoce-
rys privignis and K. truncatulus ericae.
ACKNOWLEDGEMENTS
Many thanks are due to H. Günther, Ingelheim,
West Germany, M. Josifov, Sofia, Bulgaria, I. M.
Kerzhner, Leningrad, USSR and Chr. Rieger, Niir-
tingen, West Germany for their valuable com-
ments on earlier drafts of the list, to J. P. Duffels
(Zoologisch Museum, Amsterdam), P. H. van
Doesburg and J. van Tol (Rijksmuseum van Na-
tuurlijke Historie, Leiden), the late R. H. Cobben,
G. J. Rozeboom and J. G. M. Cuppen (Landbouw-
universiteit, Wageningen), the late G. Stobbe
(Fries Natuurhistorisch Museum, Leeuwarden),
Mrs. G. Dingemans-Bakels (Natuurhistorisch Mu-
seum, Maastricht), P. J. den Boer and T. H. P. van
Huizen (Biologisch Station, Wijster), S. A. Ulen-
berg (Plantenziektenkundige Dienst, Wagenin-
gen), B. Krebs (Delta Institute, Yerseke), H. Mo-
sterdijk (Technologische Dienst Zeeuwse
Waterschappen, Terneuzen), B. van Aartsen, J.
Beernink, K. den Bieman, Mrs. C. Dolleman, G. S.
Dutmer, H. Edzes, W. H. Gravestein, Th. Heijer-
man, D. J. Hermes, R. Ph. Jansen, G. Morseld, N.
Nieser, E. J. van Nieukerken, P. Poot, F M. Smit,
A.P. J. A. Teunissen, H. Vallenduuk, H. W. van der
Wolf and J. H. Woudstra for permission to include
their collections and/or for their collecting efforts,
to N. Nieser, Tiel, for his permission to include
data from the files of the Dutch section of the
European Invertebrate Survey, to E. J. van Nieu-
kerken (Rijksmuseum van Natuurlijke Historie,
Leiden) for supplying material for preparing dis-
tribution maps and to George Thomas, South-
hampton, England, for correcting the english text.
Special thanks are due to J. H. Woudstra, Zaandam,
who introduced me to the study of Heteroptera.
His stimulating cooperation was of invaluable help
for the realization of this list.
ADDENDUM
After finishing the manuscript, two additional
species were recorded from The Netherlands:
218a. Psilophorus pusillus Reuter, f. nov. spec.
Records. - © N-Holland: Hilversum, 8.ix.1946,
W. H. Gravestein (ZMA, CB). ® N-Brabant:
Deurne, Liessel, 9.ix.1987, 1 9, B. Aukema.
429a. Melanocoryphus albomaculatus Goeze, f.
nov. spec.
Records. - © Limburg: Maastricht, 5.iii.1989, 1
6, P. Poot.
Both records will be published in more detail
elsewhere.
420. Berytinus clavipes. Scale: 1 mm.
AUKEMA:
REFERENCES
This section gives a complete list of works containing
records on Dutch Heteroptera, published since 1950. Pu-
blications, not referred to in the text, are marked with an
asterisk.
Andersen, N. M. & S. Gaun, 1974. Fortegnelse over Dan-
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Arnoud, Br., 1958. Wantsen uit Zuid-Limburg. - Natuur-
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Aukema, B., 1976. Anthocoris amplicollis Horvath, 1893
en Tingis crispata (Herrich-Schäffer, 1839), twee He-
teroptera nieuw voor de Nederlandse fauna. - Ento-
mologische Berichten, Amsterdam 36: 103-104.
Aukema, B., 1976a. Voorkomen en levenswijze van Oxy-
carenus modestus (Fallen, 1829) (Heteroptera, Lyga-
eidae). - Entomologische Berichten, Amsterdam 36:
161-163.
Aukema, B., 1976b. De Nederlandse soorten van het
genus Aneurus Curtis, 1825 (Heteroptera, Aradidae).
— Publikaties natuurhistorisch Genootschap Limburg
26: 17-22.
Aukema, B., 1981. A survey of the Dutch species of the
subgenus Hy/opsallus of Psallus (Hemiptera-Hetero-
ptera, Miridae). — Tijdschrift voor Entomologie 124:
1-36.
Aukema, B., 1983. De Invertebratenfauna van de Zuid-
limburgse Kalkgraslanden. Wantsen (Hemiptera, He-
teroptera). — Natuurhistorisch Maandblad 72: 129-
135.
Aukema, B., 1986. Psallus (Hylopsallus) assimilis Stichel,
1956 en P. (H.) pseudoplatani Reichling, 1984, twee
Miriden nieuw voor de Nederlandse fauna (Hetero-
ptera: Miridae, Phylinae). - Entomologische Berich-
ten, Amsterdam 46: 117-119.
Aukema, B., 1986a. Drymus pumilio Puton, 1877, een
nieuwe Nederlandse wants (Heteroptera: Lygaeidae).
- Entomologische Berichten, Amsterdam 46: 133-
136.
Aukema, B., 1988. Conostethus venustus fauna nov. spec.
en C. roseus weer in Nederland waargenomen (Hete-
roptera: Miridae). - Entomologische Berichten, Am-
sterdam 48: 1-7.
Aukema, B., 1988a. Orsillus depressus (Heteroptera: Ly-
gaeidae) nieuw voor Nederland en België. - Entomo-
logische Berichten, Amsterdam 48: 181-183.
Aukema, B., 1989. Pallus confusus en Psallus mollis in
Nederland (Heteroptera: Miridae). - Entomologische
Berichten, Amsterdam 49: 15-17.
Aukema, B. & M. Alderweireldt, 1989. Lamproplax picea
in Nederland en België (Heteroptera, Lygaeidae). —
Entomologisch Berichten, Amsterdam 49, in press.
Aukema, B. & J. H. Woudstra, 1985. Nieuwe vondsten
van Ortholomus punctipennis (Herrich-Schäffer,
1838) in Nederland en België (Heteroptera: Lygaei-
dae). — Entomologische Berichten, Amsterdam 45:
119-120.
Blöte, H. C., 1950. Wantsen, cicaden en bladvlooien, ver-
zameld in 1949. Natuurhistorisch Maandblad 39: 18-
20.
Checklist Hemiptera-Heteroptera 89
Blöte, H. C., 1950a. Over Graphosoma lineatum (L.), var.
italicum (Muell.). - Natuurhistorisch Maandblad 39:
113-115.
*Blöte, H. C., 1951. Nogmaals Graphosoma lineatum L.
var. italicum (Muell.). - Entomologische Berichten,
Amsterdam 13: 290-292.
Blöte, H. C., 1954. Wantsen, Cicaden en bladvlooien,
verzameld in 1950. - Natuurhistorisch Maandblad 43:
83-85.
Bozdechovä, J., 1973. Diagnostische Merkmale der Arten
Trigonotylus ruficornis und T. coelestialium. — Folia
Musei Rerum Naturalium Bohemiae Occidentalis,
Zoologica 3: 1-17.
Brakman, P.J., 1951. Oncotylus viridiflavus Goeze (Hem.-
Het.), een opvallende nieuwe Miride voor de Neder-
landse Fauna. - Entomologische Berichten, Amster-
dam 13: 199-202.
Brakman, P. J., 1952. Zeeuwse wantsenvangsten I. - En-
tomologische Berichten, Amsterdam 14: 129-130.
Brakman, P. J., 1960. Zeeuwse Wantsenvangsten II. —
Entomologische Berichten, Amsterdam 20: 21-25.
Carvalho, J. C. M., 1957. A catalogue of the Miridae of the
world. I. Subfamilies Cylapinae, Deraeocorinae, Bry-
ocorinae. — Arquivos do Museu Nacional, Rio de Ja-
neiro 44: 1-158.
Carvalho, J. C. M., 1958. A catalogue of the Miridae of the
world. IL. Subfamily Phylinae. - Arquivos do Museu
Nacional, Rio de Janeiro 45: 1-216.
Carvalho, J. C. M., 1958a. A catalogue of the Miridae of
the world. III. Subfamily Orthotylinae. - Arquivos do
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Carvalho, J. C. M., 1959. A catalogue of the Miridae of the
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Carvalho, J. C. M., 1960. A catalogue of the Miridae of the
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Cobben, R. H., 1946. Wantsennieuws uit Midden Lim-
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Cobben, R. H., 1948. Vier wantsensoorten nieuw voor de
Nederlandse fauna. - Natuurhistorisch Maandblad 37:
50-51.
*Cobben, R. H., 1948. Wantsennieuws uit Midden-Lim-
burg. - Natuurhistorisch Maandblad 37: 81-82.
Cobben, R. H., 1950. Hemiptera Heteroptera: soorten uit
Limburg, nieuw voor de Nederlandse fauna. I. - Na-
tuurhistorisch Maandblad 39: 83-84.
Cobben, R. H., 1951. Hemiptera Heteroptera: soorten uit
Limburg, nieuw voor de Nederlandse fauna. II - Na-
tuurhistorisch Maandblad 40: 52-53.
Cobben, R. H., 1953. Bemerkungen zur Lebensweise ei-
niger Holländischen Wanzen (Hemiptera-Heteropte-
ra). - Tijdschrift voor Entomologie 96: 169-198.
*Cobben, R. H., 1956. Verdere literatuurgegevens over
wantsenlarven (Hem. Heteroptera). - Entomolo-
gische Berichten, Amsterdam 16: 207-214.
Cobben, R. H., 1956a. Metatropis rufescens H. S. en
enkele opmerkingen over de overige Neididae (Hem.
Heteroptera). - Natuurhistorisch Maandblad 45: 7-
13:
Cobben, R. H., 1957. Beitrag zur Kenntnis der Uferwan-
zen (Hem. Het. Fam. Saldidae). - Entomologische
Berichten, Amsterdam 17: 245-257.
90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Cobben, R. H., 1958. Einige bemerkenswerten Wanzen-
funde aus den Niederlanden. (Hem.-Heteroptera). —
Natuurhistorisch Maandblad 47: 15-21.
Cobben, R. H., 1958a. Biotaxonomische Einzelheiten
über Niederländische Wanzen (Hemiptera, He-
teroptera). — Tijdschrift voor Entomologie 101:
1-46.
Cobben, R. H., 1959. Notes on the classification of Sal-
didae with the description of a new species from
Spain. - Zoölogische Mededelingen, Leiden 36: 303-
316.
Cobben, R. H., 1960. De eerste vondsten in Nederland
van een met Sedum en een met Carex geassocieerde
wants (Heteroptera: Miridae). - Entomologische Be-
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Cobben, R. H. & Br. Arnoud, 1969. Anthocoridae van
Viscum, Buxus en Pinus in Nederland (Heteroptera).
— Publikaties Natuurhistorisch Genootschap Limburg
19: 5-16.
“Cobben, R. H., 1979. Macropterie bij oppervlaktewant-
sen. — Entomologische Berichten, Amsterdam 39:
120-121.
Cobben, R. H., 1987. Aradus signaticornis in Nederland,
met opmerkingen over enkele andere met Pinus ge-
associeerde Hemiptera (Heteroptera: Aradidae;
Homoptera: Cercopidae). - Entomologische Berich-
ten, Amsterdam, 47: 33-38.
Coulianos, C.-C. & F. Ossiannilsson, 1976. Catalogus In-
sectorum Sueciae. VII. Hemiptera-Heteroptera. 2nd
Ed. - Entomologisk Tidskrift 97: 135-173.
Cuppen, H. P. P.J., 1978. Het voorkomen van macroptere
vormen van enkele oppervlaktewantsen in Nederland
(Heteroptera) met enige opmerkingen over hun oe-
cologie. - Entomologische Berichten, Amsterdam 38:
181-184.
Cuppen, J. G. M., 1988. Sigara 1actans nieuw voor Neder-
land (Heteroptera: Corixidae). - Entomologische Be-
richten, Amsterdam 48: 94-96.
*Dinther, J. B. M. van, 1953. Les punaises du murier
sauvage. — Tijdschrift voor Entomologie 96: 199-217.
Duffels, J. P., 1962. Vondsten van Sigara selecta Fieb.
(Hemiptera Heteroptera) in Nederland. - Entomolo-
gische Berichten, Amsterdam 22: 59.
*Duffels, J. P., 1962a. Brakwaterwantsen. - Zeepaard 22:
27-30.
*Drosopoulos, S., 1977. Biosystematic studies on the Mz-
ellerianella complex (Delphacidae, Homoptera Au-
chenorrhyncha). — Mededelingen Landbouwhoge-
school Wageningen 77(14): 22-28.
Elov, E. S. & I. M. Kerzhner, 1977. Bugs of the families
Anthocoridae, Cimicidae and Microphysidae (Hetero-
ptera) of the Mongolian People's Republic (in Rus-
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*Gravestein, W. H., 1951. Over Graphosoma italicum
(Muell.). - Entomologische Berichten, Amsterdam
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Gravestein, W. H., 195 la. Over verscheidene voor Neder-
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Tijdschrift voor Entomologie 94, verslag winterver-
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Gravestein, W. H., 195 1b. Hemiptera heteroptera: Nog
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AUKEMA:
Gravestein, W. H., 1955. Oecologische gegevens omtrent
Nabis boops Schioedte (Hemiptera Heteroptera),
Entomologische Berichten, Amsterdam 15: 395-397.
Gravestein, W. H., 1956. Nabis pseudoferus Remane,
faunae nova species (Hemiptera Heteroptera); met
een proeve van de determinatie mogelijkheden der
Reduviolus-groep (Fam. Nabidae). - Entomologische
Berichten, Amsterdam 16: 151-154.
Gravestein, W. H., 1959. Piesma salsolae (Beck 1867)
faunae nova species (Hemipt., Heteropt.). - Entomo-
logische Berichten, Amsterdam 19: 200-201.
*Gravestein, W. H., 1960. Een op Terschelling waargeno-
men merkwaardige overgang van waardplant door de
schildwants Chlorochroa juniperina L. - Levende Na-
tuur 63: 85-88.
Gravestein, W. H., 1963. Calocoris schmidti Fieb., een
nieuwe wants voor ons land (Hem. Het.). - Entomo-
logische Berichten, Amsterdam 23: 231.
*Heerdt, P. E van & W. Bongers, 1967. A biocoenological
investigation of salt marshes on the south coast of the
Isle of Terschelling. - Tijdschrift voor Entomologie
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“Heerdt, P. F. van & M. F Mörzer Bruyns, 1960. A bio-
cenological investigation in the yellow dune region of
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225-275.
Higler, L. W. G., 1962. Microvelia umbricola Wroblewski,
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*Higler, L. W. G., 1964. De Hierdense Beek in gevaar. —
Levende Natuur 67: 279-282.
*Higler, L. W. G., 1967. Some notes on the distribution
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Netherlands (Hemiptera-Heteroptera). - Beaufortia:
14: 87-92.
*Higler, L. W. G., 1967a. Hydrobiologisch onderzoek van
de macrofauna in de Grote Peel bij Ospel (gem. Ne-
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*Higler, L. W. G., 1969. Terschelling als milieu voor
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Josifov, M., 1961. Ueber drei in Bulgarien anzutreffende
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Josifov, M., 1986. Verzeichnis der von der Balkanhalbin-
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Checklist Hemiptera-Heteroptera 91
Kerzhner, I. M., 1969. Zur Nomenklatur einiger europäi-
schen Blindwanzen (Heteroptera, Miridae). — Acta
Entomologica Musei Nationalis Pragae 38: 141-145.
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Heteroptera from the far east of the USSR (in Rus-
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BD:
Kerzhner, I. M., 1984. New and little known Heteroptera
from Mongolia and adjacent regions of the USSR. IV.
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Received: 10 November 1988
Revised manuscript accepted: 13 February 1989
94
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
128. Deraeocoris flavilinea, larva V. Scale: 1 mm.
AUKEMA: Checklist Hemiptera-Heteroptera 95
INDEX an aus zul (AGT) scocsdarecodoauadaccucc0c000d 28
All insect taxa are listed, synonyms in italics. Species annulatus, Dicyphus (211) BEE i ie tei ag 17
numbers are given in brackets. annulatus, RihyNnoOcCO SIG Or) ROERO E 4, 25,73
annulicosnerd O5) Pen 3 16),
abietis, Eremocoris (462) .................ls) LE DONS LORO OE TOI 3
abietinGastrodes (465) 2452-42 en en CUT 47, annulipesh (26) has saad rca eee 13, 45
alken (AOD) eso sE AS eRe 4,28 antennatus, Megalonotus (496) .............. 29, 78
alucillom (S34). RE ao eed Aas he 3282 antennatus, MeratocorisiG46)-....--- see 22
ACIDA ora dte TEE TRE a ete ere ERS ae I Anthocondaene RE RE 23, 87
ARD ooo E n OTTO 2° WAMENOCOHIMACH ERIH NRE aoe a AT EE 23
AGAMATOSOTIA ss 5.3.5.6 IT 34, JAN thOcOrinikyaryaty Re TOTO IO E. 23
INGAMEMOSOMAtICaC may ret RTE NT OT da eo a CITE 23
PACANTNOSOMATINAER ee ee 34 Ain tilloco INSEE LITRI 28
PACE GO DS ee Pay PAPhANUSE oa icone ZONE ne CURE 4, 30
ÉCOMPOLONE Le e i ATTI QI RIVA PLREIOChEI ACRI 9, 87
PNG Tia USE I de A RAR 28) Aphelocheiru stan IEEE OOO 9
ANITA (S55) n E BAD ELO ITN 2
AGULCANOUIAEUS GIS) ccqsqonacemasecaneer anc 40, 81 INPOCHE MM MUS secures III 15
INGE TOCOBS 0.0 oda ERA 20% PABolygust. 3. ernie IALIA II 19
adenoeanpi (25 nnen EEE artan t 82 /apterustHalticus (219) nnee 17
ADERS « 05 TI TRIO 32 APtERUS IMAC (292) EC Eee Ce Ce 4, 22
AASPErsusI@ O2) A ce. GS Oe Vapterusy Pyrrhocorisi 012) ERRE NE eee 30
AR à 060 oro NERE PCE eae EE ee 334 ADI SEE LT EME ce II 22,67
ACTORS so a A RON 29) Aquariusys III On eee 2, 10
Anes (DIE) ero I II DINI (NEUES POTERE. 25, 87
email CRD) PER i IT 3720, radiale rna nen ears ete en ete RN RAS 25
PNESGhyMCELU Spee on IONE 3 Aradoidear u. a acts Neo INTO 25
aestwallie (DE). semens aleen Ro PROS ee SATA ST ie ores tee Von RR cee regu ase tere ee 25
ACUNOS à orn LE RR A COTTI IT NE HOT GTA en (IE) emo OR RO 15
ans, Calodori (GO MN eg 20, 61 ATCCOCOGISARI Wace Ace ek CLE OGL CE ETEN 2,8
ana, Corbis, (Iie ¢ ona see cee nie ou bien maid as Sie pare naniuse(©09) ERE 30
affinis, Scolopostethus (469) ................... DSE are nico lan GAD PRE TI 11, 40
ess (S13) na ve EE SOMOMMPATENOCOLIS ARE 31
AOC ooo ot III 200 APE NCAA) CIRO ST 10
NOIA IMAP n RR Ra IAD ARR cina PERLE CCE NE 34
ADICINEHI SIMONE ers 197 VARCRENCINACRA NECES 27
A] DIQU SL IAN vre OA 1A WAS CIOdE a LARE IAT RE 16
albipennis (SI) ca eo cecum ee ee VS Te IT NE ARE EERE 86
alboacuminatus (494)... a vern. ADO NAS PER hl ace ede MANN PR EC Mae Y eee 2, 39
albolineatust@0) ETRE EEE sacha: 16, 53 assimilis (A CI) RAEE RE PNR ORE ERO 15, 49, 87
albomaculatus(@20a) Lee 88 aterCapsu KCA IS 21
DORE (S30) oa do beeen masons Sona As ola 5) atersDufouriellusi (SII) E Te 4,24
aliens (CSA) 3. 85 sus ra INT, 65 aterAGEOcorisn AAI RO RT 27,75,87
VANO GAD MRAZ ts NITTO ZA terr MUSO eee OR PET ae 5, 32, 83
ANIDSOWOTIWS EEE ob OER ee 13 atomarius UO) ere TINO 19, 57
NICOLA OWS) Wee STAN ZIA CACAO MUSE TTT 15
ADESERISPRRE RA RIOT ILA ZI atrata ASD) ek RE EO 4
alpinus, Acompocoris (382) ................. 22569) “atratusy ABS) IE MEN E ARCS NE 4, 29, 78
alpinus, Plagiognathus (183) ................ 15,51 AUIECSCEMSh( 44.0) ee CRETE TE 4, 27,75
AVA ACRI ANRA USCEIA (OO) do doo aan 32, 83
PNY LU SEM INTE Saven ius Alb) 25, 74
ambiguus (ISD) soan EE LE ie haar IS. baccanum (GE): EEEN LE)
ATTINTO ER A Rc CARO O ASS baerensprungu(AUM)r EEN 24, 72
ambulanst260)e ee LOS GAMA Salis (270) a O A 19, 57
ampliema (105) EEE RE ATI 2 Bathys0en CERCA RE ARE EEE 31
amyolkeollhe GONE nenne. 22408 1888 becker BI) SE Ne 20, 61
AMOR d'os TO IE BAW ABEOSUS LETTIERI ENT A NRA LR 29
AAC 3 5.0 à lore CRE eee CS een pe Un a 20 WBErycidaer ER III eee Di eo dde 26, 87
Ameide A CU Ba otis 25%, IBERYLINAC ye RR IE Ee RN 26
Amu 35.0 ob Ed RE n 25 ABEryUDUS IT 4, 26
Annes A0) Cougs dann ER ou8 6 18 BOTY EU SiS ACLI inact CLI AA aes 4
angus ticolllisy(488) ar. EEE PE EEE RR DITA ENE EE 15)
96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
biclavatus (GO) ER EEEN E NRE CE 20, 61
bicolor OSB) Hy en EN EE LIL 32
BICUS PIS OST) Mn ren ND NINE ern 23
BIOS ae ae 34
bidentata (405) EE D>, 118)
Bieuttatus SS RE 32
Bilineatu ng (AID) re 3, 18
bilineata QIN SIIT RR APE 3
binotatus Ol 21
bipunctata (AMD en ee EREN 13
Bleplakido preti sr rr 18
BISSONE 27
Lovemant oe eee RAE 3
bohemannigG92) eee RE N 3 6, 52
CORANO OO 2
bonsdorthi(S) er pate tte oa ee ee DU
Pop BN eert pee ERE 4, 22, 67
BOthy Mo tus eleva stee MA 13
Brachyarthrumn. soi. sce I LR 14
Brachyceredicn acct neuen trae ee eee ee 17
Brachypelta. cake ea AE eee 5
Brachysteles: stekene ne det EA: 24
Brachysturar. cara EE 21
brevicollis 45) EN 1, 14, 47
Drewipenmisn(Ad3) MENEN EE 28
Bresis I GO) A a RAD A se 23, 67
Brunmeusi(45 IAA N NEE 28
BEVOCOLIMACR ER IRE OOO 13
BEYOCOLINIERI IT EE 13
BEV OCO ESTE A ende EE 13
bueno PITTI RO LGE 10, 39
butler GU ONREIN ene eae 23, 68
BIORITMI 3
caelestialiumW GA) RR 2226586
EIERN) a a i O eee 34
album ee LIA 11, 40
ealcaratag SAD) AI eee ds DI
chere OS) WER 4
CAIO ORE ee aE ee ae 31
Calloni ae Ro RO 8
CUOCO EL REIT 20
CALYPLONOLUGINA SN II RR ET 4
CampestrismEyctocoris BS) Per 4,24
campestrisgOrthopsk 2 19
Camptobrochis me EEE Sy, 3)
Gamptotelusmnne cs RI 76
Camptozyoum ee EE ECRIRE EN OUT Cee ee 20
@ampylommaret et RC NE 16
Campyloneura ee een enen EET 7
Campylosteitaree RR NRE 11
COMPOSANTE oe TR 3
CAPITAINE A LINO l
Cap rae (Oily LT ONT 2,10
CAPSOLESTE re TN 4,21
(CRÉÉ ee nee de mn Le Role ca Dai 21
capucina OO) PER NVN CPE 12, 43
Cardiasterhini Pere eN 24
GCardiastéthuS PE ese sy eacss dc I 24
CARAVAN SR RT er 12
CARIGISH (25.8) MOLLE INR ees 3, 18, 56
Cannata ZA GAlyPpral(CO RR 11
Carim Ata VA GELLO pish(355) RR TO TT Dil
Carpotoris see NEA 33
Castanea! (li) manner EERE 8
Catoplatusi.. cia PI ER E 12
CAVILLOMS: ne nd NA PR eee D
Ceraleptusi... i EEE DI
Ceratocombus:… 21221 ter 4 eee eee 7
cervinus (274) un eren 19, 57
Charagochilus:..... .. 0. ARR 20
Chartostirta.a ii ROTA 11
ChartosCistini.. SL EE ER ul
Ghilacis) ot I RR ee 27
Ghiloxanthinde nnen 10
Ghiloxanthus 22 NERO 10
chiragra (497) Re NME ere 29
Chlamydatus wa. 22... RE Re 15, 16
ehlorızans (228): nn or 2 ee 17
Chlorochroavan earn ven ve 33, 85
ChOLOSOMA torn ner EEN 31
Chorosomatini..... verd mi EREN 31
ChEysanthemil(1S9) ITE 15
ciliata (LOW) in En 12
G@iMeX tb. ss) dhe east daca CE 24
Cimicidae ar vorden ae 24, 87
Eimicoidea, nn neren EEE 22
CIMICOIdESUIC) re Loacoobacouegoe 2,8
Cimicomorphar PACE il
EINELA ((B2)) ee EEE 11
éinerea (3) vun teen I EN 7
CINNAMOMEUS (MUL) oo os ncoeoccesuocousacosce 25
cinnamopterus) (2/15) PA 17
clavatus (IG) LS UD OO ee 17
Glaviculus((G4i) eee eee IT Dj
clampes 420) EN 26, 88
Glivinem ann 13
Elosterotomus.. nn. EET 20
COCCINCANQIINN re en ee ee eee 3
COCCINEUS (ZINN 2: 2c EEE 3,18
COCKSIL (83). sua 11
coleoptratanCymatian( PETE 7
coleoptrata, Myrmedobia (118) ................. 13
coleoptratus, Ceratocombus (1) .................. 7
columbariusi(BON EEE 24, 72
Compsidolomi. ee 15
concinna (2 Mr ERE 2,8
concolor (252): u... atie EEn 18
confususmAnthocorisi(3 TT 23
COLLAND UN CO) 4,76
confusus Pilophorus (21) RE 17
contusus, Psa1lus (107) SE EEE 3, 15, 49
Comiortodes 1... RI eee 15
Conostethusi.. lira NE 14
Constrictus(206) ROLLE O REC eee 16, 53
contaminatusi( 280 19
contractus (AO) 28, 77
CREATED) EE RSS CL 650000000600 3, 12
Coranusih..:.- aen ARR RE 25
cordiger (127) ons annen a8 RE 13
Coreidaë:. Lurie EE 30, 87
Côreinae: kiss Abe EEn 30
Foresta ee RES 5, 30
Coriaceus (225) anneer een ee 3 117
Cbriomeris.. Lis ever etn IN RER 31
AUKEMA:
(CO d'a RA RT RARE 8
Core N TIA n 7,87
Cotes à Ina 7
Conbroteen. 615 ER a eae ihe 7
COS oo es N ATI 5, 21
(COnNOICARPRERNE RE RER He i ote 26
cotials (402) RE nee 25, 73
COR (154) oo ee PIE 14
costalis 30) 45000 N o cer AZIO
COS ALAN (NUN) olde dee TE 12, 43
crassicornis, Criocoris (197) ................. 16, 52
crassicornis, Stictopleurus ....................... 1
crassipes, Berytinus (423) ...................... 26
crassipes, Phymata (410) ................ 25, 73, 87
Gremnocephalusmreyes eu. 16
CASE 65.0. vr er sts cat aed ble, aoe eee BGs 16
GriSpatay ClO) EEN 12, 43
GRUGIOCUSE (233) o I 3,18
GUlICIFORIMISI(AVANE ens some 0 24
CUBRONS 0/0 0 ee a SET, ERO SA 2
GUESIFANSESCIOCONISUOPA) nn 33
GursitanswXylocoris (391). EE 24
GHRIOS (576) +4 IRR BE 34
Gralnidlae s à 302 TINI 32, 87
Oren PR MERE nn 32
CVANUSERERE SI Ale ore Ee 392
eylindricoliseeg er. eene eerde VAN
Ellecormseggogis ee le JU ZIA
GUOGOTES vus ee à
Grade or sers oo N ET 7
VIALI MACRO EIN 7
CHE. 2: n Di
CHE ole oon en 27
CYHPROSTELNUS MERE ne 34
Crodatons RER RO RE 18
COTON à 0.6.6 88 Be IA 3
chiant (SA) eee CP ee 5,31
Caron (DAO) ss eae aoe ee Ee Ee One 5
decolo EN LE 3, 14
dos (ANNE ce co 28
denticulatUSO 20) I I 31
demtnpesn GLA Papers ne emo or 8
depressus Aradus (413). 0. 25
depressus, Orsillus (436) ................. 26, 74, 88
IDYSPASO GOR Ro oc 13
IDEARC ROO |, on Ro 13
IDEFACOCOLISERERE nnt 3, 13
Deele on weeet 12
desentusWON ONB Rn ME 30, 79
chapman (245) Osea een asa. 3, 18, 55
IDG TOME Sogn as TE 7
DiChtOOSCYLUSME loos TIE. 20
DWicranocephaluspyer naa see ea eal EE TEE 4
DGA 85 brb RI O 3, 12
IDES OMCs PRE: 3,112
Dieyphinaeser OLI RTRT 16
DICYBRUS so A A e TI IR 16
dilatatus O8) MARE UE 29
AMARE 21
CRIS UC (NÉE RERO IN Eee 3, 49
diminutus Kirschbaum (172) ................... 49
DipsOcoroidear ras ES TER ee) i MOR eRe Vi
Checklist Hemiptera-Heteroptera Om
DIPSOCOLI AEREO NË VAS
Dipsocoromorphalas er 7
dispar OMD POLI 30
dissimilist(400) NEE 24,72
distincte (PO) eer ah A WE AR 8
distingendustelen Ene AMER 1
distinpguenda (ll) PEER PER PET Er 13, 45
CitOMOIdES(CA SERRE CR OA DW)
Dolichonabis ne ae 22,67
dolobratai(49) Er 4,22
A010 Data SA) cd 00 ivo 4
Dölycorisı Me TI TRE EERE 33
dominus VIET SO 34, 86
AORTA RS LTN ee RE TT 5,83
DIVANI: Be APO RA ELE eure 28
Dy Ms Faved A seen AAA 28
DEYyOPIMOCORISE TERRE E ERO OR RO 18
AUDIUSTA LERNER RE edet 1
Dufourie lis nio ero ORE eno 24
AdUMEtOtU MIMO 12, 45
dumosan(O 79) AAT ANTI OO II 34, 86
GUIMNOSUSE(OV) ek eC ER ee 86
EEN) RE RI Nee) en 12
Elasmostethus On 34
Elasmuchar as RL 34
Elasmuchinacn tees ne IS POE 34
Blatophiluse ce cata serre tat eee 23
elegantula, Chartoscirta (84) ................ 11, 41
elésantula Loricula (116) seen ene 13
elongata (340) aren ER pe 4,21
Emblethis% IT 30
ÉMESINAER ER ROL CN IONE 24
EMPICOLISEIM IATA 4,24
CNCIVISION N 2958717
ENOPIOPSRA ri ee 30
CpI LOCI RI 16
EQUESTRI 4, 26, 74
Eremocorisi: i MELO LES 28
ericaeiKleidocerys (439) LE 2, 27, 75, 88
ecicaeyNysius! AZZ) re TIE 26
éricetorum Nabisn(GOl) ere re eee 23
ericetorum Orthotylus (290) EA 18
ELEFANT 16
EEraticasNOtostiran( SAONE 4
erraticus Eremocoris BOD) nn enten 4
CIVIDNOCEDEAU ACETO ee: 53
EUAttusk A TR tr RTE 15
Eurycolpus ee eenden ene ATO AEL 1
Eurydematrt og eee ee ter one RE 34
Bury caster ane II 32
EurymEerOCOLis ha 16
IEUSAT COTES Rae ee isc steve Ree Nt oe 5
evanescens MORIRE 16, 51
CHILI) e EN: 3
IE VSALCORIS me cia a ciel geen dame CE UE DOS
fabric Cato platus(10) LOR EE E 12
TabriCn Amctocon1sa 2) ER EE 2
fabricityEysarcorisy OO) PSE CEE EEE 559
falleniMPS Alu SI QICS) RENNEN 15
jallentsPseudophloeus (SZ) er EEE PEER CPR 5
falleninSigaral(S RON 8
fallen Atenocoris, O2 REIN NT Hall
98 * TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
ASCTRENTA (BI) lodo odi desde 24,71
CO ARI 1
VETRO AIR EINE 1,54
fecaupataWEla simuch'at(054) eee ener 34
ferrugatanlkeptopternan(55,0) RER 4,22
JORGE, METS (BIO) 2 86 soccoudesdacooo ne 4
onde (EO) PROTEO CICCIO 4
LELUSH SOL) REN rr ETEN 23
FE DERIN OSD) nr ee Orr ECCO 34
RiehErOGa PSUSLAL ARIOSTO AE 197,
ESA ZIE er e 4 13
tlavellus: (169) EEE 15
flAVCOISRENRYGOPUSPRPRRE RE E 1
flaveolus, Fieberocapsus (229) ............. D 7, SA
VVLVICON NO A2) re CD PES RER 5
HAVICOLNIS (0 A2) Meen ASIAN 5, 22
Mavil ine NA 2, 13, 45, 87, 94
Havinery.ist(2A hetero cen nite mee cece Nene erent wee ae 18
favo maculatuisi(252) Pe 18
Havomarsinatusg OON A DD
tlavoquadrimaculatusy (231) Pr 3, 18
flayosparsusi(2)4) hewn. ve. RIETI 18
HO AGE CO) re eats Re a 12
fORMICELO LUTZ .osanncdeocosscucsudeoe 24,71
fossarumi IRON RI 8
fina GUI COL SA (ABS) Ey O RE OA 4, 29
LU CICO AU IA won boa RER RSD 11, 40
rallienmens (ID) oo ane aaa desc ose dou 29
fulieinosaNDictyonotan( GB) 12
fuliginosa, Odontoscelis (546) … 32
fulvicollise (DI) EEE 3, 18
IERI (IBC) ve 15,51
fulvomaculatush OA) BREE RNA E RE 20
furcata Miesovellia (AS) PER 9
VANCALAMINO LONECLAA (EB) NN 2
IMSGESICANS (QED) edo O A degne 18, 55, 88
fUSCIOLS ENER 2 Sil
MSCS OMG (HG) sc200000000000500000000000 33, 86
Lala ctinus (GO) eee 600 ERI EN 24
GAlEALUMEE Reano OI CO 12
PAU INA EME re M EEEN 83
lement (B72) e 6e oc an0sadencuc 4, 23
Ballanum (GUD) ora sen oan OE TE á
GampsoCcOLis I ee REEN 26
(Gastrodes yyy rt REC EE 28
pemellatist(@04) en 195697
SEMINAR A a CR A ee A RA 1
Fans (SI) occorsi vado Ay ET 7
Peniculatisn(435) MERE RR 29
sensae (235) ococccnccdntcovcosccsnoocs Ay IG, SS
GEOCOLIMACALIANTI MU RER TN INI RE 27
(GeOCOni sey com La O RS RENE 27
AD TOW: (Oooo aac LIO 2
GEOLOMU NEN EE D OEE l
BETA (II) 0 .0.00000 000000080000 0000.806.00 13
Berm ais (IND RNA e AMOR ARE DOSI 8
(GELLAR ITA A RANE 10, 87
Gerrinae WKE er I ARI VARI EN 10
Gerre a AS ek eee 2,10
Gerroidean ER RA OR RT EE 10
Gerromorphatrer we I I IN 9
BIDON) ee 10
AN MeT DANS 4
gimmerthali GAP 4, 21, 65, 87
Glaenocorisa =: ass 2 EE 7
glandicolo rl GAD 27)
Blauca(39).... nr EE EEEN 9
Globicepst. sn Gos chen en EEE 18
globuliferi(212) rr 17
GNAbOCONUS SLI 2 En 5
Gonianotini „un. tar stre EEE 30
Gontanotus. . „rs. Ar En 30
GonoeEruús.s ann set TY TI 30
gothicussAlloeotomuss( 154) ree 13
gothieusCapsode (911) Er All, G2
gracilenta (49) ndr rees Se ee 10
gracilissyAcalyptay ST 11
gracilis, Myrmecoris (261) ............... 19, 56
gracilis lemnostethusi G65) PAP eee eee 23
Graphosomar.sass ner note EEE 33
Graptopeltus’: 3523 233 0 PRO 29
Brisea IGO) O PO 34
griseola. arn RIE E 1
priseusiConostethusi(158) Pen 3, 14
griseus) EmmblethistO 03) meeer 30, 79
grossipes (466) 5 acc aan Sa ee 4, 28
prylloides(446))\2 7 1272 Re DI
guttula( 352) aren oe EEN 22, 67
oyllenhal CITIES 3
evillenthalitit(295) En 3, 20
Hadrodemus:... i 2245005 EMI 20
haematodest(70) es 3.15)
haemorrhoidalel O8) 34
Haldosalda=... Serre EEE 3
Elalicorimar tener dite RENEE 8
Hallodapini . 32.4 353- 5: 202 SR eee 16
Hallodapus.. ans ned RA 16
Flalosaldan inns drents nn 11
Halticocorini sr neen EEE 17
Halticus. ar iN 17
hamatus RNA 28, 77
Marpactor iure ARI eee 73
Harpactorinae.. 22... oss I NET 25
Harpocera nas vann age N 14
Hebridae : asa rai 9, 87
Hebrotdea sanar zor 000 PRE 9
ÉTÉDEUS iI 9
Hebrusella so. vene er 9
Dellensi (2A)... 2 2.0. EEE 2
hellensii (24). ben iet Se ne 2,8, 36
helveticus: (431). Sanat eee ee 4, 26
FIESPELOCOLIXA RR RR RR ERE 2,8
HFeterocordy]us:: 0% ae Ao RS 18
Heterogaster : i... net EEEN 27
Fleterogastrinae Ret NS 27
Héterorius ss 2.4... 28.300 ERETTO 23
Heterotoma':. anssen 18
iero glyph1Ca CERERE ERRE RE 2
Himacerusi u. CAN EEE 22 67
hiesutulus.. noe 2.2.2 bene 1
hirticornisi(421)te Ss EEE 26
hicrundinist(697) ee 24, 72
NISTHIONICUS (220) EN ORE 3. 17
Rolcostethust tr ANNE 5, 33}
AUKEMA:
holosericeus, Polymerus (293)............... 20, 60
holosericeus, Tropistethus (455) .............. 28, 76
holsata (597) ali RE de 4,21
bolsasna (SD) ee SONA MOT 4
TO OdEN USER TEN I INTE. 3
EIoplomachusgr.e ter tn een ee 14
I NORETLOER. EP bete NHL 83
numeri (S43) e er. 32,83
mali Dieyia (IIS) as nee 23 012.
baumalMonanthia (Qi De ee en en 3
Invallioms (527) ee anna 31,81
Bberomennts. etn INR TA 10
EIVCIOMELEIA ARR RO bee 9, 87
Fipclraomennicea 10 atteso ao denn bad sas Ban aoe 9
Bbopaillles RS nun ee 15
bpostand (SHS) Eee eden aoe 5
VOS MON oa ae anda ae BEES En 5, 31, 82
Eboceoecs RSR 2. 17
facrans (62) LR MER ee 8, 36, 87
Ibolocorts . vossen RE ia AD Iata: 16
MOCO ful à à 0 0 eee ed n cr 8
armor (Gs see Bn OE LL cao 41
taken (SS) TREE 20
means (GZ) rel 21
intermedia Rubiconia (560)..............he+s 33, 84
intermedius, Dichrooscytus (296) ............. 20, 60
INTEHSCINCHUISIO SIE I N 34
inimeacus (317) ON old E 21,61
ineusus (BP) MR EEE TES 22, 65
ia (69) RER NER 33
TEA GUN USER gens 4,73
IS CIO CONS EE TRO EN 28
ISCHROdENUS EE PE na te ee. 27
lschinorhymchinaemepy en semi oo) | Anes ence 27
SCHON NI ILE INNATO 4
ISOTMELO PINA CRE ES DD
ISOINCLO PUSH RI RN RE Faire et Eat 22
ballen (SS2N e e 5
allan seed: 34
juniperiaGonoeerussDl Oh. annen en 30, 81
junipesi@Ehytoconispee I I I 1
jumijserima SO): ES ER Re ER Cu 33
Kalama sove 12
lt (DN ee ee RE 3
kalkalt (272) sr 3,19
Illetas ree tee NN weer ws See 27
Rina (SSO)S rte Ro 33
Sten OCOLISP AIEN Ce ee 21
lastre (0) ARRE 10
ete GLIA a AS Te SI Auto 3
Ten (Mi Re i 212.
hevican (DI) ri ee en ee 4,21
VA CA au SS 4
laevis 060), 5, 25, 74
PATMPIO PARERE OOC REI 28
AS la Can that ee mie den ee to 12
FAST OS OMS SIENS MERE ER ALL 29
laterali ANGEL KONE 209)
lateraliseidalosaldag(G9) ERE Re Dali
lateralis, Sigata SD) an. erhal 2,8
lawicalllis (GBS) Er RE EEE 23, 69
VOD MIA (DEN) o's POOR VOI 4
Checklist Hemiptera-Heteroptera 99
IUS OOC CIR ES I 4, 28, 76
lectulariuss( SOS) NEER 24
Me S MOUS I RARI RICA 32
lÉPIdus CL) ER RC ee PET 15
IEptocerusn OS OPETE 18, 55
We ptopodoideara ar. ae TAGS 10
Beptopodo1morphay ERE 10
Me Pto Pte NA AA Er ese RE Be 4, 22
leucocephalusi(222) i BERE SCORE 17,54
limbatus, A NthOCOLS (072) RAP ee 23
limbatus Ey20cOnNS (282) PE PAPE SE 1959
limbatusSNabicula GS8) rn. ee 4,22
limbosuUsKO2 EEE 55.32
[imitata iSigara (CORRO eer re 8
limitatum, Brachyarthrum (153)......... 2, 14, 49, 87
Ei MO MAIS): AO ENI 22, 67
LIMAOPOUS EE RAPEN 10
linearis, Megaloceroea (341) .................... 4
linearissRanatran RE 7
lineatum, Graphosoma (552) .............. 5,33, 84
lineatus®ENabDICUl CIME EE 4, 22
Lineal usNiy S125 AEREO 00.8.0 0010 TINA A 4
lineatus, Teratocoris (348) ..................... 65
INCOME TRO een eee 5,32, 83
lineolatus, Adelphocoris (309) .................. 20
lineolatus, Camptotelust 2... toi nee 4,76
lim aen (Eene Accs ery RO SE EE 2,8
MAPPA NB e Diletta 2
TIOCOLIS lms sere Ae I CE DEN TEEN 19
Biorhyssusiays re IRR ee 31
TITOCOLISI er syste le a RT E RETRO 18
Itotalise(l 82) VARENNA RR ZO
littoralis (CORRE RR RI DE 11
Cura Us (OZONO 33
IVI LUSAZIA EAD RENNEN TETE ENNE 31
Ron eee 26
HARAS TO RE dada ne 8, 36, 87
GT ENTRE SE TE REI 21
Iongirostris (OGC) II EAN EEE PER 282307
BO Pus: x. sig ae ARTE 4,14
lo riculannens Sr AEN SII)
IACOLUM (2 SD EE En 19
hive GYAD EEE 32
LUUD TESA DBs NS NE RATA NT 2
lundi OO ER 4, 29, 78
larida Bam eran ASI ENTI 4
luridus, Pachybrachius (484) ............... 4, 29, 78
luridussBsallusi (TD) RNN 15
luridus, sStronoylocorisn 225) kerr EEE 17
lugiduslroiluss OE 34
IO IO ici toe 9, 37
luteico NS (220) ERP RER are En 1759
lutescens 24 nee Te 13
[Ey C'OCOLIMACLIA III EC LL COM PER eee 24
EY CLOEOLN eel e eren ARN 24
MEY CEO CO LIS tree Er lede ae 24
Pe CLOCONUS Ce acu Sige ep hee ad AR NAY AEN Te AEROS 4
By caeidaenic AZOTO eZ ONS
By PACINAC ILA OR LIA DCE 26
Ee A 26
Ey POCO LIS RSA ER E N 3, 19
ILS OUSE Gres aah REI N)
100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
IymEEus (AS) FARE: Dane 4,29
stavi MN COS) BERRETTO ENT 3, 20, 60
MEGA CEI TIE 3
MACCEVE CN IRINA TRE IT RIT 32
Meerodem aten N NO EI 30
IMacrolo PUS ANI ATRIA 16
Maeropartusr Ae Rhene ere TA 26
Macrosaldulatee M AI 11
Mäcrotylu Meen OI I 14
maenlaragNotonectal(Al mame ne E PISO I 9
acida Pesa (CMI) eres voto od oe av sos Bals o ord ore 4
maculatumyeresimay (DIMORE 4,25
maculatusiGaleatusi(97)) PRA E 12, 43, 87
maculatus INAUCORS OT) N e O TE 9, 37
maculatus Rhopalus SI) EPP ER E NE EEE" 31
IMAC KE LR DLE ROTTE 19
IMACMICOLMISE (ILS) IS), Sil
MARTE sn 4, 22
MMAIUSCLIGH SSO) NOR RI 4
fmnmsaulitis (HK) PEET 4, 23
MAlaCOCOLI SMELT ET 17
MUZE nt A PE 15
marginalis, Micracanthia (81)................ 11, 41
marginalis, Orthotylus (244) ................... 18
marginata, Acalypta (88) ................... 11, 42
MaLeinatuss Coreus DIS) OR aie 5, 30
mora ns (QOD) acocoosovecosckoousouscvso0s 60
IMArgiNepuncratusi (5,05) EREMO Se 30
maritimum, Monosynamma (193)............... 16
MAGI MUSABCEOSUSIALO) RIE OO SONO 29
Manitimuseley cust (20>) PE 19), 37
NIEREN (227) oo neo ano ad bom ao ao 5 2, 17, 54, 88
MUONI II o EER 32
MECOM maere Re 19
med SRM) ES EDEN 30, 79
megacephalus CAT) PAR Di UD
Megacoelumina sene ln ORE 20
MEDA OCCORRA TI 21
Mesalocoleaste: to ESRI RAR 14
MELA ONORE SO es ace eres 29
MEZZO NOISE TOR ae ee ie 4, 29
melanocephalus, Cymus (443) ............... Di TS
melanocephalus, Eusarcoris (559) ................ 5
melanocephalus,Phylus (SS) Eee 14
Melanoconyphuseee RR ea 88
melanoscelan Ar 11, 40
Melanotsichus sE 18
mers (CN ste sbsadnudedadd os dede e 25
MM OPLOST TIRO O By DI
Mesopsallus PRE ETRE 15
MESOvel er IO I on 9
MESovellda RARE EE RE RAI 9, 87
MEesovello dea Por RIT 9
Metacanthinaen LO EROE 26
RAIDS NOTE CRE DO RIOT 4
MEtarropisk heen ene De NOTE 26
MELO PORRE RO ZI U re ee ee 27
Mieracanthian EE A EN rt PAR 11
MIGFONECTI RIE TITTI IRR U
Micronectin ae ER 7
MENO WANED ois, pooh III IO 3
Micro PAYS RE TE 13, 87
GO (DIO) oddio 30
Mierosigara sd x 305 Wieren 8
Micro yncrinid ar eee eee 3
MiGrotoma i tester EEE 4
Microvelian una O 10
Mieroveliinaë. osn CODE 10
TK (GWA eg rn 23, 68, 69
minor Berytinus (422) eee ee 26, 74
minorsBlesiocori nen 1959
MINMANSEN) ian CSR 4
minutissima, Micronecta (6)............... 7, 25, 39
minutissima lea (AS) re EE 9
MINULUSA(D ET) sene EEEN 4, 23
Migidae at age sane RE 13, 87
Miridius gross tata GE 21
VIMTTONS (S32) candida 31
Mirinae®. Jenn art sE 19
Mitininesnainve nnn anr 19
Mir is atomen ee 4,20
MIRMIEOIdESI SPA) ER eee 4,22
Miroidea.. parse erheen EE 13
modestusn (449) En DID
moesta (lO ie res ANNEE RER 8
mollienlus WADE 14
mollis:(172) tr ca EER 15, 49
Monalocoris.....- su ee ene 13
Monanthia irr ent EEE 3
monereaffin( 255) oen ae EE 18
MOonosynamma EEE 3, 16
montivagusy (424) ee eee 26, 74
mOroNsaldan eee eee 11, 40
MOnioySehirus! OAI) EE 32, 83
muellerii(68)f rt en 11, 40
TMUSCH(SQ) sic os aen na ace CEE 11, 42
mútabilisn 22) err de od RARE 3
Myodochini. tri enne ee 29
Myremecorissnn 32122 11 IONE 19
Myrinedobia:. Annen en 13
Myrmus nent oaks mr ran 31
Nabieula a... sten. AE 22, 67
Nabidaes 84 ceo. vin dee sr a 22, 87
Nabinaen... avn oa beeren A E 22
Nabinidenrs 3... 0.02 es ee le 90 0.6.0 à à 0 0 0 22, 67
Na OS Eee sds = Die 210.05 d 010 8 01e 4, 23, 67
AE OC) A crei OT 10
Hassatuss(2ASN.. eu 18
DASU SIAE O E eee 14, 47
Näucoridaers i.e bene En 8, 87
Naucorinae nnn EE 8
INAUEORIS:: actin: et eea An 9
Naucoroideat} se. tc. OI RE 8
nébulosais same. ann ne er A EEE EEEN 1
nebulosus Ol 29
NOTE nes de aan Ss See 26
nemorallis(675) EEE EEEN 23
nemorums(B76) an a 23
NEO lASIOTLOPIS A EEE 12
Neolygus maan Garson son sense GEE 19
N eomecom mann TEE 18
Neopachylopsi ti tutt ACNE 18
Neottiglossa: ta sce se a IR IRAN 33
Nepa. une eee ee RIONE 7
AUKEMA:
INGE se 7,87
INISBIMAE Lies Ate TT OI Ba AS 7
NERO dE ARE AIR 7
IN PO O Ep AE ne PE 7
miser (DA) o RR Re CSS 423
20200 BOA) es 4
miemiconninNCarpoconts OO. ee nee 86
nigricornis, Elatophilus (368) ............. 23, 68, 88
MORNAGO REI eae ene ee Se 12, 42
DE, ACH (OLE) 32, 83
mignitamPolymerusi(294)\ ennen nende LE 20, 60
EOL TN UO EERE EPE 152
migcolineataySigara(2))) Le de ZG
nigrolineata, Trigonosoma (552) ................ 84
MORVELICU MRO OE 20
INOEONEEEAMEN RN mr ee ct TIRI 9)
IN@IOMEGHEACS à 05 na Hen OH Been Goo e Ad ahd 9,87
INOLOMECINACH ee CO Eee 9
INGtOMECLOIG CAR ee 9
INIOLO SATA EI rn ce RITIRI N 21
MOTS INR eere RI RARI 62
nubiluswbathysolen\(23))- . an... ne 31,81
nubilusssMacrolophusi(205) ona 16
nubilussPemtrechusa(491) RENE 29
INVSÎOG 0 ET a ee CINA EE: 26
OBERE) ER er IR ea IR ES 12
ObliquanNotonecta (43) 2. ee Mk 259
OUTED, Cane (CAD a nn SO 4,75
obsentellusk lim ee ZD
obsoletumt Olien E NE 16,92
oalromelas I i 3
Odonto Paste COPPA II III USE: 10
Odontoscelistn TT ORI 32
DECIA CUS ER IA INT AANG 24
oleragsn (STD se IO en 34
Clivacsus (120) srate e oo 15
ONCOcH ARR ee I 12
OMG UNE IO AI 14
OCEAN OO EPA 3
ouala QD) ER ec er 11,41
Online RER ER nt Ae 75
OP 5e Le SO TE 23
OLSENE IN RI ANA 4,23
ODA LUBIANA 5, 86
Oria. ARR OS RR a: 26
Crsilhs … ae 26
Croce EE Ce ee: 17
ormhoedilla MO» i ovens e e ano 11
@rtholomuse TEA re CCS 26
OLCHONOISERE EEN ITA TI 14
CLEO PSP eo INI By Als)
OO NACRE RÉ 17
ORVIETO 17
OLCOWIUSB I E Sean 2a ee 3, 18
Oxyeareninae ree AIAR 27
OXRYCATEN USER NRB nen IA ION 27
pabulinusW(2 PEREN 19
Pachybrachiussn nnn E OO E 29
Bachiy CO leusy A nnen ER PEER RE 7
Bachyimeruse a IAT 79
Pachytomela fe RE AIRONE 17
PAPER ete on bord 14
Checklist Hemiptera-Heteroptera 101
allons CO) ot On pont ane ZIO
balidiconnss 20 3
pallidipennis 75
pallidust CONT en ee RO ERE ER 16
PallipeS CAT) rata e IE 11, 40
Balomenan a. Lo aaa IE 33
paludums/Aquarius (63) RIEN PE 10
paludumsdieratocorisn( SAT) EEn Ce 22, 65
palusteissPolymerusy(290) EERE 20
palustrissSaldulan(@/8) Wrs NE 1, 11, 40
JHA ae EE TONE 4
Pantiliust branen EE 21
PINZE. ROIO CO TER FER 8
PACACOLIXAMR RITRAE 8
PATALIOPICEpPS RER nn AUTRES 18
parallel 224), SP E ARRE 17
ParapsalluS es cute mener IR EL 15
parumpunctatus (29) Erne CEE MERE NE 31
PAGVICOLMSICI IEEE ENT CE 24, 71
PAM OLE AO ME ES DN 12
parvulus, Atractotomus (180) ................ SN
parvulus EX YIOCOTIS EE EEE I IE 1
Paykullun@a AI): se eener OI 3,14
Pedest rss (ABZ) eo TEE 4
REntatom a EER 33
Bentatomidae ee 33,87
Pentatominae tn 33
Pentatomordea en 32
Pentatomonpha yaaa ers II 25
Peri ZA MRP O N >)
Perittechus RI AO 29
perplexis (218) ZORRO 17
PELLIS 2) RAM ne RI 15
Personatus (AWG va 25
Bhimoder aa TERN 32
phoeniceus(492) aa ase ee 29
BhOEnicocoris NER 15
Phyll dea i er ER AE AR EN 15
Bhylinae A cso AA EE SEN 14
Phys ANNE EE 14
PROS ME thes ed SO e SAI 14
Bhymatak en ni e 25
Bhymatınae sa. TE 25
Physatocheilan IAA 2
PISA sense ee EEE 3
Phy COCOLIS qr ee LE TINTI 21
Pia NA CE) IR I CRT 4, 28,77
PICeUs (A608). ar ES pence EE 4,77
DEIN USA CORTI, CARITAS TE
Pipe SD I) NME Geter sera tuyere RATE D5 92
Picromerust ee NI III A 34
pietus (AD aster eenen AE 28
Piesmar Ma ee en eae 25
Biesmatidae ee TE EN 25, 87
Piesmatoidea re or neee TN aE TOE 25
Biezodorusi a. Re LE 33
PEZOSECLD US: nn NE EE 4
pilicornis\Calocons1(B00) Fe ES ITEM 20, 61
pilicosnissDrymusn (AS) ee ET EE 28, 76
Pier IO OO NZ?
Bilophotin EST NE EL RO RA 17
Philo} NOG A EE RE RARE RER 197.
102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
POS AQLOZ) EEE Mere a dina o's Sip er 12, 43
pilosellaN eects ce xe RIE AR 11
PilosussAnthocorisn O7) PERRE 23, 68
pilosussBothynotus) (23) ee ee 13, 45
PilosussChiloxanchusi(G>) REE 10
pilosussMevalocoleusi(/43) EERE: 14
pilosus, Scoloposthetus (472) ................ 28,77
Binalitusar I OENE RIN AE 3, 19
DIR AS En OSD ATTICO 3
PINETE 3, 14
UATE ODIO en DI RI 3
pin, Amoer (ID) ERRORE 21
pini, Rhyparochromus (493) ................... 29
pinicolan( CO) NA III 33, 85
Binocapsus. may LIO EE 18
PA NENAEUS EDS ME II IA 1
IPiGNOSOMUSin ats VITO EEN 30
DEDEN AN EO) ENE 72
pipistrel AO) ER Ae RO RE DS TD
BIL COAT ASSO TAO I AIR 85
PiCRA MAIA e nee RN RZ AA 19
Pithanus gien PO 19
PityopsalWSi TI 15
PIC gNathusyiswor ssc ee 15
Plan co: SKI) AE ISO 3, 18, 55
platychela 02) ee ee 3, 12, 42
Macon CP) ciocca code 3, 42
EN Te es DR NO EUR RE 9
PIEDEUS AR TE Ne ARRET 28
BIE AEN 9, 87
PIETER O 9
BJESIOCOE STR E IE E 19
EIESIORE MAREA 14
EICSIOVCNA ERE ERRE RR PE 10
BA thisimie rennen EE 27
Plm ESSA Heel: 28
BIOCIOmeEnI se 78
PIGTATIO A ET ee AO OO. NAT 4
poagrious (GOS) RR REI ot a’ 28, 77
Rodopinde ATI cess eu OT 33
ROMO PS ts MI ee 33
Roeciloseytusi rat en Seed od eather aoe ee 3, 20
PO LOPES III IAA 15
Bolymerus nennen I II 3,20
POPULIMEYLOCOLIS SI) PRE 2 10, 59)
POpulPhytoconiSO 20) APP SERE E EE 21
DOME) A INA cae eae es 7,36
pates GOD) 0e 0000006 0000 shout don 29
PERSON RER AR PPT de 1,8
Pras1naN CD) MO ni done nata eco 33
PEASIMUSH (ZAG) IMC AN ES RN 18
(RAKES (AO) PER 19,97
Privienisi(437) BARRA DAT ANS
POULE ST A CNIT 1
pPLopinguay (lO) PETE DU
Brostem ma I ROTA
Brostemmatinae EE O VE A
Prostemmatinies er EE ARTE?
BEOXVIOCONIS NOT 24
PAUSE RN 115)
psammaecolorm (S43) DD
pselapmifoemisn di ee scopes 13
Pseudoferusg(& 6A) ERRE eee ee 23, 67
pseudograndisi(475) Pe 2, 28, 77
PSEULOlOXO PSE RE ET RETE 18
Piendophlaeus SIA ARE 81
RONDES eos sseseaaccecodoaooscosesee 5,81
Pseudophloinde eee EEE 31
pseudoplatanin(d!63) mre EEEN 15, 49, 87
Pseudoveriniconixal Sd En 8
pteridis( 122) neer ee eek See EN 13
Pterotmetus : 242 2. iu ace O 30
Puberulus (474). nn PR RE 28
pulchella GOOR 2, 24, 71, 88
pulchellus BA)... 2 E 22
pulica usi) RECARE ARESE eee ee 15
pullus(U89)… aren SIA ARRE 16
Ppumilion 99) Eee EEE EEEN 28, 76, 83
punctatal (UG) torens ren ATTORE 2,8
punctatonervosus) O35)... PE ae eee 32, 82
PUNCTATUS (DE) occ aaacacacc0nnccc0cccceacaccs 34
PUNECIPENNIST(35) PARRA 4, 26, 74
punctipes, Gampsocoris (427) ................ 4, 26
punctipes, Oncotylus (136) … 14, 46
punctulatus, Deraeocoris (125) ............... 13, 45
punctulatus Geotomus .:. EE 1
puUEpUreipenmis) (6.68)! rr 33
pusilla, Neottielossa GSM) PEER DI
pusillus Elebrus) (417) ane eee OR 9
pusillus, Pilophorus (218a)..................... 88
pusillus, Plinthisus (454) ................... 28, 76
pusillus, Temnostethus (367) ................... 23
PYCNODLETNA Nn 3
pygmaea, Microvelia (54) .....:........... 1, 10, 39
pygmaeus, Acompocoris (383) .................. 23
pygmaeus; My tthus (2) lee ae 14, 47
Pygolampis. encore 0 25
PYHOIDES) uns sn re ET 1
Byrrhocoridae tt: ER EEE 30, 87
Pyrrchocanis..........2.:22.2. 200 eee 30
quadrata Bıesana (IEEE 4
quadratum, Piesma (418) ................. 4, 25, 74
quadratus, Xanthochilus (495) ................ 4, 29
quadrimaculata partim (109) ................. 3, 45
quad rsmacúlataypartims (LOWER ee 43
quadripunctatus, Adelphocoris (308) ............. 20
quadripunctatus, Calocoris (302) ................ 61
quadrivirgatus(@15) CREER 21, 61
quercus (HOO). are O EEN 15
Raglius. Mise ss ee OI IE 29
Ranatian nanne veder RI IT 7
Ranatrinae. iui EE 7
reclairen(267) ri N IE 3, 20, 59
TEGUCORNISH GADE 4, 21, 65
Reduviidae nnen OO OE 24, 87
Reduviinae i ET 25
Reduvius. anc oaren MRS TI TT 25
Reduvoidea venne TEE 24
resedae (438). D NN IE 4,27
reticulata, IMicrovelia (>). 0)
reticulata, dungis) (ON) SR eee 12, 43
REtrOCOEIXA oars amer RC CI 8
reuters Notomectan(44) a as oe ee ence 9,37
TEUTERIEHYIEOCOLISLS2 ERO SR 21
AUKEMA:
REUSEL PENN ee 17
Riel Ie eer orne de 20
RURACOSMAUIUS foo soldat ORE 34
Fhasanicola (OS), EN er hier nadia 19
RNA PRASSI 1
RIDIRE O EE EE 4,73
dnrococenae (OS) ROLO RE 12
TRONCA (NM): 0 san ve 5
Khombeusko ERRE ee ANNE 5, 30
Rho palid ae au ea n RE 31,87
Rhopallimas.. oe sau RE RO RIS ERA TERN. 31
RFO PAT nd ree oee en 31
RINO PALI SRNR pees ne BAE DAL
RIVM O CO LINRA 25
Rinyparocehrominael. nn. e 29
Rhy parochrominiye so ee een An 29
RUG SAROC OTIS oe cee Ed 4, 29
WOERD + een On de 12
moflamdel (SOD) as onee eo To I dens bh 4, 30
HOSEOMACUIATUS PO) A Beieren en eee 20
rose (199) sos den OR A 3, 16
roseusqConostethusi((139) iena 14, 46
none alas MO O 3
rocnmmmal (200) PE ER EN N 3, 16
ROOTED (QO 655 SE RENO 3
mbar (130): sin er ENEN terete ee Ton 13
lem 5. 6-0: 5) n Seen a A 33
rubicundusn(Fallén)\(288)). .. dhe... nr. 20, 59
rubicundus sensu Reclaire (287) .................. 3
DICE (296) oa bele NE PET EN RO 18
rubens (279) N Rai gabe 19,5%
rufescens, Metatropis (428) ................. 26, 74
rufescens, Hallodapus (202) ............... 3.216,55
niescensmBachycoleus (ON rarr ae... 202,35
GUACEPRCNPST RE EN Er RE 9
OUBICORMISMC AS) ee 22 (656
rotinons (MAN) cea doses a EE daran 3,14
DENTS (CO) Wedde ARIE en Eko IE oe 20
KUÉIPESFACGOMPUSIAMO) NL nt AE PRE 28
rufipesPentatoma (07) rte ee er 33
rmifosanteliamas (EL PE AE 3,10
AUS à 0: 01070 5 RE dae 1
muarcollig (278) va A deere Henst sne Eeen ne Te 19
CURSUS. (HOA) AE RN 23
gueulipennisN OON) anr. e a 19
FUSTICUSE CO) ONT 29
El (LEO) eta os RE OI 128 7
Sabulleni (AAA) EE EE 27
sabulicola, Megalonotus (500) .… 1, 29, 79
sabulicola, Monosynamma (194) ............. 16, 52
gabullosns GSA). 6 REE 4, 29
SAND SO) ER ewer ews obese ale EE 8
Salla so sah NATI TIA 11
Salilicae: sr Sees RARE REI 10, 87
Saldimae HER 11
Cale LINEE AR AE eet i SUR Ob 11
GA ARRE iat oie REA ADONE, LIRE 10
Sellenmigr ss Sen A RE 16
SACE ZO EER ee 3,15
Alco 3, 15, 49
SANTIS E IO E EI: 3
SAISON ETC) Heten da se O 25
Checklist Hemiptera-Heteroptera 103
Saltatons lal tics: (22/1) PEER ER 17552
saltator, Orthocephalus (226) ............. 17454555
om) Ar 00 LL EE 11, 41
Saltitam ssl OOMEN en ren 16
SANPUINIPES 2 en M EE 1
SarOthamnil Gio) re 23
Sa DI enor re 10, 37
SANTE MO AS) NE 22, 65
raie (CAO) Le usb PORGERE 26, 74, 87
Scaplag UN renem EE 5, 30
SCAHADAEDITE SO 4) ee TE 32
schillincin33) aera SCR ae 31
Schimid tin ONE 20, 61
chnesderi (OD) vreden teen MRI 2
SCHOZIUONNER LO COLI AIN 7,35
SCIOCORIS er a en LE 33
SCOLO PIER ILA KANE 24
Scolo poscelistty uan are FEN: EAN 24
Scolopostethuser. OI TRA En 28
SCOLIEAN MON NEO EA AREN 11, 40
SCHEN ee AE E 8
scutellaris 1 ERE 13, 46
ScutellEridae ENE 32, 87
SEhIEINAE re TE AE 32
SEH EE IE 32
SC1ECtA 22) es EP rea ere 8, 36
semistriatan (AI) ne INA 8
SEMEGIONISN(AS4) Er NE ET 26
SPENDEER N See 3
Seticornis OMO TE EE 20
SID IEIEUSDI TORI NIRO NITTI AN 68
SIDAT A II UAAR 2,8
SIENALIEOLNISL A A) I 25, 73, 88
sipnoreti (Mn ATI 26
IMPERO III OA 12, 45
FIERO EN apice n'a 23, 69
singer 2A EEE 262
smreezynsictil (109) PPR 312245
Soler (GUAM ES RE I E 14, 47
Spathocera ..: saret Seen CRUE 31
sphacelatust@ 0) Pon 33, 84
sphepitormisn 23 OI II 18, 54
Sphrapistieus ITTICA OT 29
Spilosterhus RETE 4, 26
SPINACI NE 19
Stapmallisn(23) ewes TEEN 2,8
SCALO UM ONEROSO 10
StalieN NE DDO
staphiliniformsKÓ OS FERA. domes eee re 4, 30
Staphylinoides(608) ITER eee 4
Stenocephalidae ur... TO TT 30,87
STEN OGEND A UNIT REN 4
Stenodemarg,. nrs maen 21
Stenodeminiss singent ehehe NI 21
Stenopodainaer nee ceases soe cit ete 25
Stenotus nnn NICO EO 21
Stephanitisv ARI, Macey. MIRE eh ERIN 12
STRENATUS CR MNT ERP TNT ATA 3, 16
Stictopleurus MER TEASER ee 32
Striata (29) ontario RNA Qt A as: 8
ARITA OIE ee ee 3
State line (GON ys oo bob bade E 3, 20, 61
104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
SERIA NON AAA A 3, 20
SERIE DOCETAN OA) eenen 12
SELOM SY IOCOLISH Lit nre enti etn EE 17
SEVP NOCOLINI ee AE 28
S GY MO GOLISR eye AI N tia AAE 29
Subaprecush 408) PERRE DS),
A PALA ZA (IBD) co 000008055 0000H A000 RnB A000 46
SUDEUFUSOS ON AAA 31, 82
SUbSIgara ii crc ete AE 8
suleaans (332) 0006000000000 2, 21, 62, 88
SUIGICOEMISK (OS) PPE 2, 16, 52, 88
Sylvadyrnusrs tse aren EA AR 28
Sylvaricusi( A ON) rr AEN ren ene 28
VOT ra NGO eo ee ER 4, 78
DO UDP TEEN 3
ISVEOMASTUSLELLA ICONA III 30
SYSCELONO USATI IO 16
laphropeltus@sen nes re I AE 28
lemnestethus:. are ee 23
tenella, Myrmedobia (120) .........:............ 3
tenellus N\Orthotylus (247) BREE Eee 18
PRETACOCOLIS eo TROIE 22
testudinarial®>))) Sennen aoe eee oR RO 32
NETTA PIE PSE ION EEEN 23
WE Ken ES dn AEN 83
UDP DAN Doll
thomson) EAN 28
thoracica lar poceral(150) PERE REN RR 14
thoracicuse Gerrisn (Gil) PE EP EE EN eee 10
drone (I) BRERA DENE 14, 47
EDU DELTA Rn 47
Cini (BB) ee ooren eme ee 26
Dhyreccoridae TTI 32,87
IR YECOCOLISE ene er ee enge ee AN 32
ele (ZIN ere Hee ER 18
tieinensisg Gilly) PPT 20
OE (RD) TESERO 21
PELO Id ae erneer RI 11, 87
TIERRA NOI Jul
MBN S ASH Ei Weeen EN 12
Hiingordear...un- une un IRE 11
tipulariusy(A26) ea TER 26
(ira PEZONOLUS is. unse ee EEE 30
IDFICNOCALOCOLISI LA ARAN 20
CRICOLMIS (OON EEE IE 3, 12
toifasciatusM GSD) i My LL EER 13
Wig o moty lust Eeen. eon RARA en IE 22
UNI guttatuss(2 03) FERRE 16
TTD LED AGRO EEEN 4
tri pustinlatusi(260) PETE 19
tBISPINOSaU (BAO) coccoccecuggavccas00nau00 4, 21,65
PRES PINOLIA (BBO) oocccsaccceccosaccnasccagccns 4
ERISEDIALUSR OOO) Piri TO 34
IEIEOIME PAS iro I 32
(ro iu Ae O ena Clete bid OR 34
ropidoche la ran res 12
Hiropidoph [eb IA EE Dj
PICO PIMYSIUSH Nen en Hen Re RAR 26
Dropisterhusm naren ern iere en 28
trumcatulusk SO) 2, 27, 75, 88
tubereulifer > same: doc Oe eee 1
TUMICICOLNISU(23 SERRE SORA 18, 55
tunicatus!(330)) cao nne EEE 21
ty pliae(49l)bt.. an een O eee 27
Aytthus: antennen EE 3, 14
UMBRO 21
DDD TION (SB) et 39
UMPEINUS Er en thse Soe EEE Il
Unifasciatusi 29) En 20
urticae 452) bo siti 27
wacabundus) (404) PT 24
variabilisn(l64) 23.5 ve EEE 15
varlansı (7A). 2er ee EDES 15
Varipes (226) „annen A REEN 21
VAIUSMDOND 5 tre EPEN 30
Melia ni ante nd IP 10
Meliidae + eht anni En 10, 87
Meliinaers nv AE 10
VENALOT.A(DIS) ES Wk ee 81
ventralisO7A) see 1, 5, 34, 86, 87
venusta,(Sigäranranor nenten oe RT E 1
venustus, Conostethus (140) ................. 14, 47
verbasci, Campylomma (196) ................... 16
verbasci, Emblethis (504) ................... 30, 79
VENUS REN RL eer ee ee 5
VermicOrixal a tit RI RE 8
Verna (89) teen ten ie bee 5 I
Vernalis:(962) aar LCI SRE EEE 33, 84
HAN ON nenn ee 40
vicinusi(288) rare nr ern EEEN 23, 69
virens, Orthotylus (248) .................... 18, 56
VirensyStenodema (339) rn 21, 65
virescens (252). mito RE PT 18
Virgula: (213) 2 eee 17
vunıdiflayıı std) Eee 14, 46
viridinervisi(249) MRE 18
viridisslyoocorisM 282) PETER 19
viridiswNotonecta 42 9
Viridissimal(0.04) ERRO re RE E 33, 85
VISCIMANINOCOrISE (O 80) meen 23, 69
viscinHypseloecusn(2l4) ERESSE E 17,53
viscicolau(276)..Ar PRE REP EEE 19) 57
vitellinusi(1 7) PP 15
vulneratusi(292) EEn 20, 59
wagner CAPSUS D 2I) EE EEE EEE RSR 21, 62
Wagneri, Lygus (268) "FREE EEE REP EEE 2, 19,57
waeneri, Psallus (165) PP PERRET 15, 49
waltiMPachycoleus\@) FAP sane aoe RE 271,35
waltls:Preudophloeus (522) EE EEE RE 5
waltliiArenocoris (522) Eee 5, 31, 81, 87
wollastoni: ron 0e NUL En 1
woodroffein 40) RO REA 11, 2, 25, 73
NANTNOCRILUSE FEES 29
XYIOCOTINI ver reerde ORE 24
XK ylocorisif paces re 0 one EE 4, 24
Zieronâ wanten been EN 34
TWO NEW MORDELLIDAE (COLEOPTERA) FROM KASHMIR
AND SUMATRA
by
R. BATTEN
ABSTRACT
Batten, R., 1989. Two new Mordellidae (Coleoptera) from Kashmir and Sumatra. — Tijd-
schrift voor Entomologie, 132 (1): 105-107, figs. 1-8. [ISSN 0040-7496]. Published 3 July
1989.
Two species, Mordellistena leveyı from Kashmir and Calyce sumatrensis from North Sumatra,
are described as new to science.
Key words. — Mordellidae; taxonomy; oriental region.
R. Batten, Sprencklaan 3, 4333 HC Middelburg.
INTRODUCTION
Thanks to the kindness of Mr. B. Levey (London)
and Mr. K. Wellschmied (München), who both
sent me Mordellidae for identification, I am able
to describe two species as new to science.
The following abbreviations are used: l/w -
length-width ratio; py/hy — pygidium/hypopy-
gium length ratio; w/l — width/length ratio;
RMNH - Rijksmuseum van Natuurlijke Historie,
Leiden.
Formula for the ridges: 3-4.3.2.2. means: hind
tibia (without apical ridge) 3-4, first segment of
hind tarsus 3, second and third segment two ridges
each.
Mordellistena leveyi spec. nov.
(figs. 1-5)
Holotype male: Kashmir, Srinigar, 26 June 1984, leg.
B. Levey (in RMNH).
Paratypes: 5 males and 4 females, same data as holotype
(one female, labelled allotype, in RMNH, other para-
types in collection Levey and Batten).
Description. — Belongs to Mordellistena s.s.,
having each of the first four segments of the an-
tenna narrower and shorter than segment 5. It
belongs to the gemellata-group, characterised by
two short lateral ridges running more or less pa-
rallel to the apical end of the hind tibia (apart
from the apical ridge) (Batten 1980, Ermisch
1969).
Dimensions in millimeters of holotype and al-
lotype respectively: Length without pygidium 4.1
(4.5), length of elytra 2.6 (2.9), width of elytra
105
together 1.0 (1.2), pygidium 1.6 (1.1) and hyp-
opygium 0.5 (0.3).
General shape elongate, sides convexly atten-
uated posteriorly. Integument black. Pubescence
black with a reddish hue.
Head w/l 1.5. Finely punctate, hind margin
slightly convex, maxillary palp of male (fig. 1);
the same of female with anterior segment broadly
scalene, innerside rounded in middle. Eye finely
granulated, hairy, no temporal margin. Antenna
length ratio segment 1-5: 10.10.7.10.11 (fig. 2);
segment 5-10 1/w in male 1.7, in female 1.1.
Pronotum w/l 1.3, broadest in front of base,
anterior lobe protruding, basal lobe prominent,
lateral border in dorsal view convex, in lateral view
moderately so, basal angles oblique with a rounded
edge; punctures shallowly impressed. Scutellum
triangular with a rounded point.
Elytra l/w in male 2.6, in female 2.4; sides
slightly convex, attenuated from shoulders to apex;
apices separately rounded, punctures slightly as-
perate, pubescence covering integument. Pygidium
slender and conical, py/hy in male 3.2, in female
3.3. Legs in male with fore tibia almost straight,
without a fringe of hairs; apex of penultimate tar-
sal segment of fore and middle legs straight, middle
tibia shorter than tarsus, hind tibia with two ridges
parallel to the apical one. Ridges 2.2.1.0. Spurs
yellow, length ratio 1.3; length ratio of tarsal seg-
ments: anterior 2112411015; middle
35.22.16.10.15, posterior 55.40.20.15. Parameres,
apex of penis and phallobase, see figs. 3-5.
106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
4
5
Figs. 1-5. Mordellistena leveyi spec. nov. — 1, maxillary palp;
v
IS
2, antennal segments 1-5; 3, parameters; 4, apex of penis;
5, apex of phallobase. Figs. 6-8. Calyce sumatrensis spec. nov. — 6, maxillary palp; 7, left temporal margin and eye
in lateral view; 8, hind tibia with ridges. Scale bar. Figs. 1-7: 0.1 mm, fig. 8: 0.25 mm.
Discussion. — This new species is the second
entirely black species in the Mordellistena gemel-
lata-group with two yellow or yellowish red spurs
on the hind tibia. M. lindbergi Ermisch (1963)
differs from M. leveyi by the antenna with seg-
ments 5-10 almost square in the male (square in
female); by a broader pygidium (py/hy 2).
Etymology. - Named after its collector, Mr. B.
Levey.
Calyce sumatrensis spec. nov.
(figs. 6-8)
Holotype female: North Sumatra, Dolok Merangir, De-
cember 1969, 1050 m, leg. E. Diehl (in RMNH, ex coll.
K. Wellschmied).
Description. - Dimensions of holotype: Length
without pygidium 4.1 mm, length of elytra 2.9 mm,
width of elytra together 1.2 mm, pygidium 0.7 mm,
hypopygium 0.5 mm.
General shape broad. Integument of elytra yel-
BATTEN:
lowish brown, darkening posteriorly; head with
mouth parts, palps, antennal segments 1-3 and legs
yellowish red; pronotum, metasternum, metacoxa
and antennal segments 4-11 dark brown. Pubes-
cence yellow.
Head w/l 1.5, finely punctate, hind margin con-
vex, maxillary palp club-shaped, apex cup-shaped
(fig. 6). Eye finely granulated without hairs, tem-
poral margin very broad, anteriorly ending
abruptly (fig. 7). Antenna length ratio of segments
1-5: 10.10.10.11.9, segment 5-10 serrate l/w 1.25,
segment 11 elongate to oval.
Pronotum w/l 1.8, broadest just before base,
anterior lobe protruding, basal lobe slightly prom-
inent, lateral border convex in dorsal view,
abruptly narrowing apically, in lateral view slightly
convex (almost straight), basal angles oblique with
rounded edge, punctures shallowly impressed. Scu-
tellum triangular with a rounded tip.
Elytra 1/w 2.4, sides almost straight, slightly
attenuated in the posterior quarter, both apices
rounded, punctures slightly asperate; pubescence
thin, integument still visible. Pygidium abruptly
conical from middle to apex, py/hy 1.4. Legs
straight, apex of penultimate tarsal segment of fore
and middle legs deeply excised, middle tibia shorter
than tarsus, hind tibia with two longitudinal ridges
parallel to the tibia (fig. 8); ridges 2.2.1.0; tibia with
two yellow spurs almost equal in length (ca. 0.3
mm). Length ratio of tarsal segments: anterior
20.10.7.5.11, middle absent, posterior 50.23.14.20.
Discussion. - This new species has a very broad
temporal margin in a right angle, abruptly ending
at the eye, which is a generic character of Calyce.
According to Ermisch (1949-1950) the genus
Calyce is found in the tropical and subtropical re-
gions of America and Africa. C. sumatrensis is the
first species of this genus from Asia. It can imme-
diately be distinguished by the presence of two
instead of five or more ridges on the hind tibia (cf.
Ermisch 1943).
ACKNOWLEDGEMENTS
I am very grateful to Mr. B. Levey and Mr. K. Well-
schmied for making this material available for
study, to Mr. B. Brugge (Zoölogisch Museum Am-
sterdam) for material from Java and Sumatra and
to Mr. J. Krikken for valuable advise.
REFERENCES
Batten, R., 1980. Notes on the Mordellistena gemellata
group (Coleoptera, Mordellidae). - Entomologische
Berichten, Amsterdam 40: 41-45.
Ermisch, K., 1943. Die Gattung Calyce Champ. (Mor-
Mordellidae from Kashmir and Sumatra 107
dell.) nebst Beschreibung vier neuer Arten. - Entomo-
logische Blätter 39: 53-60.
Ermisch, K., 1949-1950. Gattungen der Mordelliden der
Welt. - Entomologische Blatter 45-46: 34-92.
Ermisch, K., 1963. Die Mordelliden der Insel Cypern
(Coleoptera, Heteromera, Mordellidae). — Notulae
Entomologicae 43: 49-67.
Ermisch, K., 1969. Familia Merdellidae. - Die Kafer
Mitteleuropas 8: 160-196.
Received: 19 July 1988
Accepted: 30 December 1988
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FUMASTACIDAE (ORTHOPTERA) FROM THE PROJECT
WALLACE EXPEDITION TO SULAWESI (INDONESIA)
by
R. K. BUTLIN, R. E. BLACKITH & R. M. BLACKITH
ABSTRACT
Butlin, R. K., R. E. Blackith & R. M. Blackith, 1989. Eumastacidae (Orthoptera) from the
Project Wallace Expedition to Sulawesi {Indonesia). — Tijdschrift voor Entomologie 132:
109—113, figs. 1-6, table 1. [ISSN 0040-7496]. Published 3 July 1989.
All six eumastacid species found in or near the Dumoga-Bone National Park (North Sulawesi)
belong to the subfamily Mnesicleinae. Four species, in the genera Pseudomnesicles Descamps,
Karnydia C. Bolivar, and Samartella Descamps are new to science. The male of Karnydia
celebesica Ramme, hitherto known only from the female, is described. All the species are endemic
to Sulawesi, although Samariella is represented in the Philippines.
Key words. — Mnesicleinae; taxonomy; karyotype; ecology.
Dr. R. E. Blackith, Zoology Department, Trinity College, Dublin-2, Ireland.
INTRODUCTION
Wallace (1869) comments on the general paucity
of animal species on Sulawesi, and despite inten-
sive collecting, few individual eumastacids were
found. The finding of nymphs of Karnydia gract-
lipes Ramme feeding on Selaginella ferns and of
adult Psexdomnesicles species inhabiting the Gym-
nosperm Gretum L. was recorded by Blackith
(1987). Ramme (1940) described K. gracilipes
from N. Sulawesi (Minahassa) and C. Bolivar
(1930) described K. celebesica from the same re-
gion. Three species of Pseudomnesicles were pre-
viously described from Sulawesi; P. roseosignatus
(Brunner) from Toli-Toli in the north of the island
but some 350 km west of the Dumoga-Bone Na-
tional Park within which Project Wallace operated.
P. plagiator (Ramme) and P milleri (Ramme)
were both described from the south of the island.
All three form, as Ramme (1940) notes, a close-
knit group of species, and the similarity of their
male genitalia as figured by Descamps (1974a)
confirms this view; the terminalia are the means by
which most eumastacid species can be distin-
guished.
MATERIALS AND METHODS
Nymphs and some adults were collected by
sweeping vegetation near the ground. Eumastacids
109
in rain forest appear to be mainly canopy dwellers
and collecting them involved reaching a fallen tree
within hours of the crash or picking up specimens
brought down from the canopy by fogging with an
insect-paralysing solution, organised by a team
from the British Museum (Natural History). Both
methods yielded meagre numbers of eumastacids.
One eumastacid was taken in a Malaise trap run by
the same team.
Abbreviation. — BMNH = British Museum
(Natural History).
RESULTS
Mnesicleinae
Pseudomnesicles rhodopeplus sp. n.
(fig. 1)
Holotype @: Indonesia, Sulawesi Utara, Minahassa,
Toraut Forest, 13.1.1985 (BMNH fogging team)
(BMNH). Paratypes 1 8: same data, Malaise trap on R.
Toraut bank (BMNH fogging team) (Museum Bogorien-
sis). 2 9: same data, swept on forest floor. (Blackith)
(National Museum of Ireland; BMNH).
Virtually identical with P. roseosignatus
Brunner, except for the phallic complex, whose
dorsal view is shown in fig. 1 (cf. figs 47, 49 and 54
of Descamps (1974a) for the phallic complexes of
110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
the other described species in this genus). The
shaft of the penis valves of P. rbodopeplus is at
least 4 times as long as in other species.
Dimensions. — Elytron length 11.0 mm; pro-
notum length 2.9 mm; hind femoral length 10.9
mm, width 2.8 mm.
Females almost indistinguishable from those of
P. roseosignatus figured by Ramme (1940) except
that the lateral processes of the trifid subgenital
plate extend as far back as the median one, whereas
in P. roseosignatus they are shorter.
Dimensions of female — elytron length 13.7
mm: pronotum length 4.1 mm; hind femoral
length 13.8 mm, width 3.4 mm.
A putative food-plant for this species (Gnetum
sp.) is noted by Blackith (1987). The name
rhodopeplus signifies bearing a roseate shroud.
The red flush at the base of the elytra, notable in
this genus, is much less marked on females in life.
Samariella aulaeiadventa sp. n.
(figs. 2-3)
Holotype &: Indonesia, Sulawesi Utara, Toraut Forest,
13.11.1985 (BMNH fogging team) (BMNH).
Head as in fig. 2, distal 5 antennal segments
compressed, pubescent. Scape produced into boss
forwards, with clear patch at tip. Face rugged. Ely-
tra reaching genicular region of hind legs. Outer
dorsal margin of hind tibiae with ca. 18 strongly
unequal teeth; inner margin with ca. 18 more re-
gular teeth. These margins fuse into a wide (ca. 1
mm) tooth subequal to longest tibial teeth. Termi-
nalia as in fig. 3; phallic complex relatively unscle-
rotised, penis carried at right angles to body as in
S. bakeri Descamps (cf. Descamps 1974b, fig. 68).
Dimensions. — Elytron length 9 mm; pronotal
length 2 mm; hind femoral length 8 mm.
The name aulaeiadventa describes one who de-
scends from the canopy, originally of a Roman
theatre. The two species previously described in
this genus are both endemic to the Philippines.
The only specimen of S. bakeri known is a male
whose phallic complex differs from that of S. aw-
laeiadventa. The other species (viduata Descamps)
was described from a female. The vertex in
aulaeiadventa sets it apart from the other two
known species.
Karnydia celebesica C. Bolivar, 1930
(fig. 4)
The genus Karnydia, of which K. celebesica is
the type species, is endemic to Sulawesi. It was
described from a female, and since specific distinc-
tions depend mainly on the male terminalia we
include a description of the male.
Material studied. — 1 @: Indonesia, Sulawesi Utara.
Gunung Muajat, 29.vii.1985 (BMNH fogging team)
(BMNH).
Fastigium truncated distally as in Bolivar’s de-
scription of the female. Antennae set well below
midpoint of eyes, almost as low as unpaired ocellus
(i.e. lower than in Ramme’s (Ramme 1940) figures
for K. gracilis Ramme). Antennal organ promin-
ent conical, projecting distally from 10th of 12
segments. Body fulvous, abdominal tergites 1—3
and 8— 10, scape, underside of fastigium and genae
in subocular region and teeth and dorsal carena of
hind femora fuscous. Fragae on hind femora weak.
Hind tibiae with 19 teeth (longest 0.15 mm) on
outer carena and 15 on inner (longest 0.7 mm).
Hind femora with 5 teeth on mid-dorsal carena.
Fore- and hind-tibiae clad with at least 50 golden
setae. Male terminalia as in fig. 4; subgenital plate
and its lateral lobes, seen from rear, enclosing
roughly equilateral triangle.
Dimensions. — Pronotum length 1.8 mm; hind
femoral length 10.5 mm, width 2.0 mm. Because no
material was taken zn copula we cannot be certain
that this male is K. celebesica, and even if a pair
were obtained identification of the female might be
ambiguous.
Karnydia monki sp. n.
(fig. 5)
Holotype 4: Indonesia, Sulawesi Utara, Gunung Po-
niki “Ice Station Zebra”, 16.x.1985, (K. Monk) (BMNH).
Very similar to K. celebesica except for the ge-
nitalia, the more compressed and raised dorsal ca-
rena of the abdominal tergites, and the red flush on
the postero-lateral angle of the episternum, the
proximal half of the fore- and mid-femora, and on
the distal parts of the hind legs. First two flagellar
segments pale, remainder fuscous. Fragae on outer
flanks of hind femora more strongly developed
than in K. celebesica, dorsal carena of hind femora
with 7 teeth. Hind tibiae have 19 teeth on outer
carenae (longest 0.2 mm) and 5 on inner (longest
0.6 mm).
Terminalia as in fig. 5; the subgenital plate, seen
from the rear, forms a V-shaped slot or inverted
triangle quite distinct from that of K. celebesica.
Dimensions. — Pronotum length 2.3 mm; hind
femoral length 11.5 mm, width 1.9 mm.
Karnydia laticlava sp. n.
(fig. 6)
Holotype 4: Indonesia, Sulawesi Utara, Gunung Mua-
BUTLIN, BLACKITH & BLACKITH: Eumastacidae 111
0 0.5mm
Figs. 1—6. — 1. Pseudomnesicles rhodopeplus sp.n. Holotype 8, dorsal view of phallic complex; 2. Samartella
aulaciadventa sp. n. Holotype 8, dorsal view of head; 3. Samariella aulaeiadventa sp. n. Holotype 6, dorsal view of
terminalia; 4. Karnydia celebesica C. Bolivar. Dorsal view of & terminalia; 5. Karnydia monki sp. n. Holotype g. Dorsal
view of terminalia; 6. Karnydia laticlava sp. n. Holotype @, dorsal view of terminalia.
Lowest scale bar for figs. 1, 4, 5 and 6.
112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
jat, 4.viii.1985. (R. K. Butlin) (BMNH). Paratype 9:
Indonesia, Sulawesi Utara, Edwards’ Camp, 4.viii.1985
(R. K. Butlin) (National Museum of Ireland).
Similar to K. celebesica. Antennal organ excep-
tionally thick and prominent. Flagellar segments
dark brown, scape, pedicel, and proximal fifth of
first segment white. Integument translucent, ex-
cept for opaque yellow discs underlying basiconic
sensilla. Hind tibiae with 16 teeth (longest 0.8
mm) on inner carena, 17 teeth (longest 0.1 mm) on
outer carena. Terminalia as in fig. 6, subgenital
plate trifid with 2 hooked lateral process and ar-
row-head median process.
Dimensions. — Pronotum length 2.4 mm; hind
femoral length 10.2 mm, width 2.3 mm.
Female. As for male, but mouthparts, cerci, distal
parts of ovipositor valves, subgenital plate, and
legs livid green. Lateral carenae of pronotum pink,
underlined with fuscous. Hind femora with 11
teeth on dorsal carena, hind tibiae with 19 teeth
(longest 0.8 mm) on inner carena, and 19 teeth
(longest 0.2 mm) on outer carena.
Dimensions. — Pronotum length 2.8 mm; hind
femoral length 14.6 mm, width 2.3 mm.
The name /aticlava describes the broad, distally
spatulate, cerci of the male.
Karnydia gracilipes Ramme, 1940
Nymphs and adults were collected at two loca-
lities; Gunung Mogogonipa (altitude 1000 m) and
Gunung Ambang (altitude 1200 m). Both moun-
tains have abundant ground cover of mosses and
ferns. The species was taken from several ferns
including Cyathea spp. (Cyathaceae), Dennstaedtia
sp. (Dennstaedtiaceae), Nephrolepis sp. (Nephro-
lepidaceae) and a species of Athyrium or
Diplazium (Athyriaceae). It was also taken from an
angiosperm Elatostema sp. (Urticaceae). The role
of ferns in the nutrition of tropical grasshoppers
has been documented by Rowell et al. (1983). Pa-
latability tests of these tropical plants were con-
Table 1. Feeding preferences of Karnydia gracilipes
Plant species
Gunung Mogogonipa
Palatability*
Elatostema sp. 16
Cyathea sp. 1 8
Cyathea sp. 2 0
Dennstaedtia sp. 0
Nephrolepis sp. 9
ducted in the laboratory at Base Camp using 5
nymphs of K. gracilipes for each test, which lasted
for 24 hr.
The results are shown in table 1. Nymphs of K.
gracilipes evidently prefer Elatostema but also eat
several ferns, though not Dennstaedtia. This result
accords with feeding tests by Blackith and Blackith
(1966) on Australian eumastacid grasshoppers
(Morabinae). These are generalist feeders but con-
sume several ferns, as do K. gracilipes. Rowell et
al. tested the palatability of ferns of the genera
Dennstaedtia, Cyathea and Diplazium for the Co-
sta Rican eumastacid Homeomastax dentata (Saus-
sure). Interpretation of their results is complicated
by strong specific differences within plant genera.
Diplazium expansum being highly palatable whe-
reas D. aff. herbaceum was among the least pala-
table. Cyathea multiflora and Dennstaedtia
obtusifolia are also relatively unpalatable. These
authors note that even if the grasshoppers prefer
a diet of a single plant species, the structure of rain
forests makes general feeding almost obligatory.
CARYOTYPIC STUDIES
Although not all the species taken could be exa-
mined, testis squash preparations of two of them
were made. K. gracilipes proved to have a diploid
number of 21 (2n = 20 + X) which is common in
the Eumastacidae. There are ten pairs of acrocen-
tric chromosomes graded in size with a medium
sized X. K. /aticlava, however, proved to have 2n
= 19 (18 + X) including a very large autosomal
pair, probably by fusion of two of the larger auto-
somes of the closely related gracilipes. As is com-
mon in other eumastacid subfamilies, speciation in
these Mnesicleinae seems to be accompanied by
chromosomal rearrangements (White 1977).
DISCUSSION
Our findings support the view that only one
subfamily of the Eumastacidae has reached Su-
Locality
Gunung Ambang
Palatability
20
*) Palatability is defined as the percentage of the available leaf consumed by the eumastacids, out of approximately
20 sq. cm offered to batches of 5 nymphs.
BUTLIN, BLACKITH & BLACKITH:
lawesi. Speciation in this subfamily, the Mnesiclei-
nae, follows the pattern for the family of close-knit
genera, species within genera differing little except
in the male genitalia and, where investigated, in
the caryotype. As all 9 species known from Sula-
wesi are endemic, and only one of the 3 genera has
representatives elsewhere (on the nearby Philippi-
nes) little biogeographic information can be gle-
aned from Sulawesian eumastacids.
Altough collecting from rain forest canopies is
usually difficult, the small number of adults of spe-
cies other than K. gracilipes collected by members
of the expedition is striking, amounting to roughly
one specimen per person-month of collecting, even
when aided by fogging equipment rarely available
to collectors. As collecting was spread over most of
1985, seasonal effects are unlikely to be responsi-
ble. We are forced to the conclusion that eumasta-
cids occur at very low densities in these rain forests,
possibly partly because the great diversity of host
plants may mean that acceptable hosts are them-
selves thinly dispersed.
It is hardly surprising that, under conditions
favouring sibling mating for lack of much oppor-
tunity to meet unrelated individuals, speciation in
this group involves chromosomal rearrangements.
Sibling mating has been suggested (White 1977) as
a way of allowing such rearrangements to reach
homozygosity despite the often demonstrably un-
favourable effects on the heterozygote. Virtually all
eumastacid species whose caryotype has been inve-
stigated differ cytogenetically from one another.
With the discovery of two further species of
Karnydia a sequence of changes in the structure of
the male genitalia becomes apparent. This se-
quence runs through celebesica — monki — lati-
clava — gracilipes such that the postero-lateral
processes of the 9th tergite are reduced, tergite 10
becomes less sclerotised, and the postero-lateral
processes of the subgenital plate become elongated
and contorted. The terminalia as a whole become
broader and the cerci distally more strongly sclero-
tised and spatulate from celebesica to laticlava alt-
hough gracilipes has cerci comparable with those
of celebesica. So far as we can judge, the morpho-
logical contrast between celebesica and laticlava
corresponds to the fusion of two autosomes in the
caryotype.
ACKNOWLEDGEMENTS
We are grateful to all those concerned with Pro-
ject Wallace, particularly the support teams, and to
Dr. K. Monk and members of the British Museum
(Natural History) fogging team for permission to
examine their hard-won material. RKB acknowl-
Eumastacidae 118
edges with gratitude facilities and advice provided
by Prof. G. Hewitt and financial support from the
Percy Sladen Memorial Fund. This paper is based
on material collected whilst the authors were par-
ticipants in project Wallace, sponsored by the
Royal Entomological Society of London and the
Indonesian Institute of Science (Results of Project
Wallace no. 24).
REFERENCES
Blackith, R. E., 1987. Primitive Orthoptera and primitive
plants; 124—126. — In: Evolutionary Biology of Ort-
hopteroid Insects. Ed. B. M. Baccetti, Chichester, Ellis
Horwood.
Blackith, R. E. & Blackith, R. M., 1966. The food of
morabine grasshoppers. — Australian Journal of
Zoology 14: 877—889.
Bolivar, C., 1930. Monografia de los Eumastacidos. —
Trabajo del Museo national de Ciencias naturales, se-
rie zoologia, Madrid 46: 1—380.
Descamps, M., 1974a. Diagnoses et signalisations d’Eu-
mastacoidea (Orthoptera) II Indo-Malaisie. — Bulle-
tin du Muséum national d’Histoire naturelle, 3me
série, no. 237: 1033—1053.
Descamps, M., 1974b. Les Mnesicleines des Philippines
(Orthoptera, Eumastacoidea). — Bulletin du Muséum
d'Histoire naturelle. 3me série, no. 262: 1653—1996.
Ramme, W., 1940. Beiträge zur Kenntnis der Acridien —
Fauna des indo — malayischen und benachbarter Ge-
biete (Orth.), mit besonderer Berücksichtigung der
Tiergeographie von Celebes. — Mitteilungen aus dem
Zoologischen Museum in Berlin 25: 1—243.
Rowell, C. H. F, Rowell-Rahier, M., Braker, H. E.,
Cooper-Driver, G. & Gomez, L. D. P., 1983. The pa-
latability of ferns and the ecology of two tropical forest
grasshoppers. — Biotropica 15: 207—216.
Wallace, A. R., 1869. The Malay archipelago: the land of
the orang-utan and the Bird of Paradise. — vol. I,
London, Macmillan.
White, M. J. D., 1977. Modes of speciation. — San Fran-
cisco, W. H. Freeman.
Received: 23 January1989
Accepted: 16 March 1989
114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
BOOK ANNOUNCEMENTS AND REVIEWS
Williams, D. J. & G. W. Watson, 1988. The scale insects
of the tropical South Pacific region. Part 1. The ar-
moured scale insects (Diaspididae). — CAB Interna-
tional Institute of Entomology, Wallingford: 1—290,
figs. [ISBN 0 85198 608 0 — Price including postage
in UK £ 26.95, in USA US $ 57.00, elsewhere
L 2995)
All armoured scales are sap-sucking insects and
many cause considerably damage to crops, damag-
ing fruit and reducing yields. Scale insects world-
wide number about 6,000 species, of which 2,000
are armoured scales.
This taxonomic account of the armoured scales
(Diaspididae) of the tropical South Pacific region
is the first of three parts planned for all the scale
insects (Coccoidea) of the area. Keys to 37 genera
and 124 species, including 38 new species, are pro-
vided, accompanied by detailed descriptions and
illustrations. The main purpose of the work is to
provide agricultural staff who are concerned with
pest control and quarantine inspection in the South
Pacific area with a reliable means of identification
of species. Non-specialists can also use it to identify
and study this destructive group. The area covered
is Melanesia and Polynesia, from Irian Jaya in the
west to Easter Island in the east, but Kiribati in
Micronesia is also included. The work provides a
record of the armoured scales of each territory and
of the islands within each territory; knowledge
essential for effective quarantine inspection and
for export of plant produce. Records are based on
material examined from the major world collec-
tions of Pacific scale insects and on the economic
literature. The work covers all the important pest
species of the area including Aspidiotus destructor
Signoret, Parlatoria cinerea Hadden, P. pergandii
Comstock and Unaspis citri (Comstock), and con-
cludes with a comprehensive list of the host-plants
and their associated armoured scales.
[From information of the publisher]
Williams, D. J. & G. W. Watson, 1988. The scale insects
of the tropical South Pacific. Part 2. The Mealybugs
(Pseudococcidae). — CAB International Institute of
Entomology, Wallingford: 1—261, figs. 1—95. [ISBN
0 85198 625 0 — Price including postage in UK £
22.95. in USA US $ 47.95, elsewhere £ 25.25].
The second of three parts planned for the scale
insects (Coccoidae) of the tropical South Pacific.
Keys to 37 genera and 92 species, including three
new genera and 28 new species, are provided with
detailed descriptions and illustrations. The work is
designed to provide agricultural staff with reliable
means of identification of species and to encourage
non-specialists to identify the group.
[From information of the publisher].
Brown, R. G. & I. D. Hodkinson, 1988. Taxonomy and
ecology of the jumping plant-lice of Panama (Homop-
tera: Psylloidea). — Entomonograph 9: 1—304, figs.
1—150, tables 1—3. [Published by E. J. Brill, Leiden
etc.; ISSN 0106—2808; ISBN 90 04 08893 8 — Price
136.00 Dutch guilders = appr. US $ 68.00].
The major aim of this study was to provide
accurate descriptions of all Panamian psyllid spe-
cies together with keys for their identification. The
Panamian fauna contains a majority of the known
Neotropical genera and this revision should there-
fore provide a more solid foundation for future
taxonomic and biological work on the South Amer-
ican fauna. A parallel set of aims was to examine
the evolution, host plant relationships, zoogeo-
graphy and seasonality of the Panamian psyllids.
[From the autors’ introduction].
A REPORT ON A COLLECTION OF ODONATA FROM NORTH
SULAWESI, INDONESIA
by
R. R. ASKEW, G. G. CLELAND, D. A. L. DAVIES & T. W. HARMAN
ABSTRACT
Askew, R. R., G. G. Cleland, D. A. L. Davies & T. W. Harman, 1989. A report on a collection
of Odonata from North Sulawesi, Indonesia. Tijdschrift voor Entomologie 132: 115-121, 1 fig.,
2 tables. [ISSN 0040-7496]. Published 3 July 1989.
Odonata collected in the first quarter of 1985 by two participants in Project Wallace, the Royal
Entomological Society of London Expedition to Dumoga-Bone in North Sulawesi, are listed with
observations, and distribution of the species is discussed. The dragonfly fauna is principally
Oriental in affinities, but primary habitats harbour a high incidence of endemic taxa.
Key words. - Odonata; Sulawesi; Indonesia; Distribution.
Dr. R. R. Askew, Department of Environmental Biology, University of Manchester, Manches-
ter M13 9PL, UK.
INTRODUCTION
As a prelude to his description of Celebophlebia,
Lieftinck (1936) drew attention to the fact that up
to that time no general account of the Odonata of
Celebes (now Sulawesi) had been attempted. Al-
though he clearly stated that his intention was to
produce such a work, this task remained incom-
plete at his death in 1985. An opportunity to inves-
tigate the little-known odonate fauna of the north-
ern peninsula of Sulawesi (Sulawesi Utara) was
afforded by Project Wallace, the Royal Entomolog-
ical Society of London — Indonesian Institute of
Sciences expedition in 1985. Two of us (Askew,
Harman) participated in Project Wallace in the
early months of 1985 and the material collected has
been examined by Cleland and Davies and is re-
ported upon here. Further recent collections, Lief-
tinck’s detailed notes and the bulk of the historical
material from Sulawesi are at Leiden Museum
where they are being worked on by Dr Jan van Tol
(1987a, 1987b). Additional relevant material is in
the Hope Collections (Oxford) and the British Mu-
seum of Natural History (London) and this, to-
gether with descriptions of new taxa collected on
Project Wallace, will be dealt with in future publi-
cations.
The expedition’s base camp was situated in the
Dumoga-Bone National Park at 0°33’527N,
115
123° 54’21”E, some 200 m south of the River (Sun-
gei) Toraut. Most observations were made within
2 km of base camp, a lowland area (alt. c 200 m)
that included a good diversity of dragonfly biotopes
in both the lowland primary rainforest to the north
of the Toraut and in the land cleared for agricultu-
ral purposes to the south. Records from Sulawesi
Utara from the region of Menado, Labuanagi (a
village on the north coast), Danau Mooat east of
Kotamobagu and Kolintang about 6 km north of
Malibagu on the south coast are also included, as
are a few from the islands of Sangir Besar (San-
gihe) (visited by Askew) and Ternate and Bacan
(Batjan) (visited by Harman), and from Maros in
south west Sulawesi north of Makassar (material in
Davies’ collection). Figure 1 illustrates the location
of collecting sites.
ANNOTATED LIST OF SPECIES
Taxa from the Toraut base camp area are num-
bered. Details of the three pond sites in this area
are as follows:
Riverine ponds. — Temporary rain pools at the
edge of the forest on the north bank of S. Toraut.
Base camp pond. — A small, clean pond of re-
cent formation about 5 square metres in area and
40 cm maximum depth almost surrounded by
116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
SANGIR BESAR (|)
HALMAHERA 1)
Labuanagi TERNATE
B
Danau Mooat
Q
BACANS ©
ES
SULAWESI
S. TUMPAH
Maros
S. TORAUT
Kitchen pond
0 Rs £ 5 i
400 metres È ee a — >
nt di Base Camp“.
Base camp pond *
Primary forest i
Roads & tracks -.---------- 3 Sq
Fig. 1. The principal collecting sites in Sulawesi and Halmahera, and a sketch of the Project Wallace base camp area
on Sungei Toraut.
bushy secondary vegetation but with few aquatic or CALOPTERYGIDAE
emergent plants.
Kitchen pond. — One of a series of pools in a Neurobasis kaupi kaupi Brauer (1)
linear, marshy depression with a slow through- Toraut, S. Tumpah and first tributary. Regularly
flow. Somewhat polluted by domestic effluent but observed through February and March on this fast-
supported fish. flowing river, a major tributary of S. Toraut. Males
ASKEW ET AL.:
make frequent sorties from their perches on rocks
in the river, their blue hindwings very conspicuous.
Only occasionally rest on vegetation. Females less
often seen, usually flying close to water margins.
Mostly observed where flow-rate of river reduced.
CHLOROCYPHIDAE
Libellago rufescens (Selys) subsp. nov. (2)
Toraut, S. Tumpah and first tributary. Males
seen frequently in February and March, perched
upon rocks in fast-flowing sections of the river.
Made brief sorties from perch to return after a few
seconds. Difficult to follow in flight against ripples.
No females noted.
L.r. rufescens (Selys)
S.W. Sulawesi, Maros. Two maies, one female of
the nominotypical form, collected 18.ix-9.x.1983,
in Davies’ collection.
L. xanthocyana (Selys) (3)
Toraut. Observed from 17.11.1985 through
March only at one place on S. Toraut where males
perched on rocks and jammed logs in the centre of
the broad river. Seventeen counted in an area of
about 4 square metres. No intersexual behaviour
observed but males display to one another, a flying
male hovering in front of and facing a perched
male; latter spreads wings and raises abdomen.
Only a few females seen, usually near the banks.
Rhinocypha monochroa Selys
S.W. Sulawesi, Maros. A pair in Davies’ collec-
tion, 18.ix - 9.x.1983.
R. frontalis Selys (4)
Toraut, small forest streams feeding S. Toraut
and S. Tumpah. Scarcest of the three Toraut chlo-
rocyphids and noted only on small streams where
males perched on bankside vegetation. Females not
seen.
R. tincta semitincta (Selys)
Bacan, Fort Bentang and Air Blanda. Three ma-
les, one female, 7,8.111.1985.
LESTIDAE
Lestes (Paralestes) praemorsus (Selys) (5)
Toraut, riverine pond. One male, 25.11.1985.
Odonata from North Sulawesi 117
COENAGRIONIDAE
Agriocnemis femina femina (Brauer) (6)
Toraut, Labuanagi, Bacan (also Java, Bogor).
Common at base camp pond from January to end
March and also a few on kitchen pond. In neglected
rice paddy at Labuanagi. Polymorphic with male
abdomen dorsally all black or with orange-red co-
louration on segments 8-10 and posterior third of 7
(numbers of captured specimens 7:4 respectively).
One example of an intermediate male form with
only segments 9+10 red also taken. Immature fe-
males had abdominal segments 1-6 lemon yellow,
7-10 black, whilst mature females had 1-6 bluish,
7 reddish and 8-10 ochreous blackish.
Argiocnemis rubescens Selys (7)
Toraut, base camp pond, two males in February
and one female in January. A second female taken
in the forest in March.
Ischnura senegalensis Rambur (8)
Toraut, kitchen pond (also Java, Bogor). Only
two of each sex taken.
Pseudagrion crocops Selys (9)
Toraut, Menado. At Toraut two of each sex were
collected at kitchen pond, base camp pond and in
the forest.
P. celebense Lieftinck (10)
Toraut, kitchen pond, single males on 26.ii and
6.111.1985. A female at Menado, 7.11.1985.
P. ustum Selys (11)
Toraut, one male at base camp 14.1.1985.
Ceriagrion sp. nov. Ì
Danau Mooat, one male 25.111.1985.
Ceriagrion sp. nov. 2
Bacan, Fort Bentang, a pair 8.iii.1985.
Teinobasis superba (Selys) (12)
Toraut, Ternate, Bacan. This species, with elon-
gated abdomen, was frequently seen flying, some-
times in tandem, in the lowland forest at Toraut.
It was usually in the vicinity of small forest streams
but often flew up to the canopy. Very variable in
size, two males from Toraut being very small but
otherwise resembling closely the rest of our mate-
rial.
T. helvola Lieftinck
One female in a coconut plantation near Dolo-
118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
duo, about 8 km S.E. of base camp, 1.111.1985. One
male, Bacan, Fort Bentang, 8.111.1985.
ISOSTICTIDAE
Selysioneura sp. nov.?
Sulawesi Utara, Koluntang, one female
29.1.1985. Attributed to this genus with some un-
certainty. Most representatives of the family are
confined to Australia, New Guinea or small islands
off the N.E. Australian coast, but Se/ysioneura is
known previously from Halmahera.
PROTONEURIDAE
Nososticta flavipennis (Selys) (13)
Toraut. Two pairs taken in tandem flying in the
forest close to S. Toraut, 10.i. and 21.ii.1985. The
female of the latter pair was ovipositing in a pool
connected with a deep, still section of the river.
Nososticta sp. nov. 1 (14)
Toraut, forest edge. Two males, 20.1.1985.
Nososticta sp. nov. 2 (15)
Toraut, S. Tumpah, one male, i.1985.
PLATYSTICTIDAE
Identification of the following three species
pends publication of a revision of Sulawesi Platy-
stictidae by Dr J. van Tol.
Drepanosticta sp. 1 (16)
Toraut, forested W. bank of S. Tumpah, one
female 9.iii.1985. Probably conspecific with a male
and female taken at Kolintang (19.1. and ii.1985
respectively).
Drepanosticta sp. 2 (17)
Toraut, forest edge, one female, 8.1.1985.
Protosticta sp.
S. Tumpah, altitude 900 m. Two teneral males,
20.11.1985.
AESHNIDAE
Anaciaeschna jaspidea (Burmeister)
Doloduo, 8 km S.E. of base camp. One male at
light, 22.00 h, 20.11.1985.
Heliaeschna filostyla Martin (18)
S. Tumpah. One male flying slowly downstream
near confluence with first tributary, 15.iii.1985.
Shortly before his death, M. Lieftinck, in a personal
communication to one of us (Davies), suggested
that H. filostyla should be placed in a new genus.
This possibility will be dealt with in a future publi-
cation.
Gynacantha basiguttata Selys
Bacan, Air Blanda, one female, 7.iii.1985.
G. bayadera Selys (19)
Toraut. Forest near S. Tumpah, two females,
12.1.1985 and (at rest beneath palm frond)
27.11.1985. A large, dark dragonfly, seen several
times flying low and fast through the forest and
always evading capture, may have been the male of
this species.
G. penelope Ris
Bacan, Air Blanda and Fort Bentang, one male
and two females, 7,8.111.1985.
G. rosenbergi Brauer (20)
S. Tumpah, one male, i.1985.
An anactine species was seen flying out of reach
near the base camp pond on three occasions.
CORDULIIDAE
Hemicordulia assimilis Hagen in Selys
Bacan, Air Blanda. One female, 7.iii.1985.
Macromia irina Lieftinck (21)
S. Tumpah. Two males captured in March, one
at altitude 900 m and the other near confluence
with first tributary at low altitude (c. 280 m). Three
or four observed at latter site, hawking about 2 m
above a deep pool in the river but very evasive.
LIBELLULIDAE
Tetrathemis irregularis leptoptera (Selys) (22)
Toraut. Four males caught 21.ii.1985 at pool for-
med by inlet of river at forest edge, and others seen
subsequently at this place. One male by S. Tumpah,
10.1.1985.
Nannophya pygmaea Rambur (23)
Toraut, Danau Mooat, Labuanagi, Menado. At
Toraut observed only amongst marginal vegetation
of kitchen pond.
Cratilla lineata (Brauer) (24)
Toraut, base camp. One male, 7.1.1985.
ASKEW ET AL.:
Lyriothemis cleis Brauer (25)
Toraut, base camp. One male, 24.1.1985. Larvae
of this species develop in water-filled rot-holes and
a number were found in the forest adjacent to the
base camp (Kitching 1986).
Lathrecista asiatica (Fabricius) (26)
Toraut, base camp. A female (15.11) and a male
(5.iii.1985) taken.
Agrionoptera insignis quatuornotata Brauer
(27)
Toraut. Two males at pools in partly dried-up
forest stream, 18.ii and 2.iii.1985.
Nesoxenia mysis (Selys) subsp. nov.
Bacan, on coast 15 miles N.W. of Labula, one
female, 4.iii.1985.
Diplacina militaris Ris (28)
Toraut, Sangir. Only males found at Toraut, by
S. Tumpah and small forest streams.
Diplacina sp. nov. 1 (29)
Toraut. Two males and a female captured flying
along the course of a small forest stream, 26.ii and
2.111.1985. Another female observed ovipositing
unaccompanied. One male by S. Tumpah.
Diplacina sp. nov. 2 (30)
Toraut, base camp area. One male, 20.1.1985.
Diplacina sanguinolenta van Tol (31)
Toraut. One male in forest north of river,
12.111.1985. A distinctive Diplacina with abdominal
segments 2, 3 and most of 4 blood red, the rest
black. Recently described (van Tol 1987b) from
material accumulated by Lieftinck from a range of
localities in Sulawesi and Banggai Island.
Potamarcha congener (Rambur) (32)
Toraut. Common about base camp area, at ri-
verine ponds and base camp pond.
Orthetrum glaucum (Brauer) (33)
Toraut, Menado (Mt. Lokon). At Toraut one
male taken at riverine pond and a female on S.
Tumpah.
O. chrysis Selys (34)
Toraut. Males taken in February and March
mostly on the banks of S. Toraut but occasionally
penetrating a short distance into the forest. One
found devouring a large cyclorrhaphous fly.
Odonata from North Sulawesi 119
O. sabina (Drury) (35)
Toraut, Labuanagi, Danau Mooat, Menado (also
Java, Bogor). Common at Toraut hawking amongst
low vegetation on banks of S. Toraut, at mouth of
S. Tumpah and at the very edge of the forest.
Diplacodes trivialis (Rambur) (36)
Toraut, Labuanagi, Danau Mooat, Menado. The
most abundant anisopteran at Toraut, hawking low
over the ground on the base camp area and espe-
cially along paths. A female was observed oviposi-
ting unaccompanied in base camp pond.
Crocothemis servilia (Drury) (37)
Toraut. A single female in field of maize,
11.11.1985.
Neurothemis stigmatizans manadensis
(Boisduval) (38)
Toraut, Labuanagi, Danau Mooat, Menado, San-
gir. This conspicuous red-winged insect was com-
mon but with a very aggregated distribution in
rank secondary vegetation near standing water.
Males spent much time perching on vegetation,
often only 1-2 m apart. Abundant in rice paddies
at Labuanagi.
N. fluctuans (Fabricius) (39)
Toraut. Much scarcer than its congener above
and represented in our collections by just one of
each sex taken on the base camp area in March.
Neurothemis sp. nov. (40)
Toraut, Bacan. Allied to the above. Four females
captured, two on the base camp area, one by S.
Tumpah and one on Bacan (Fort Bentang), in Ja-
nuary, February and March.
Trithemis festiva (Rambur) (41)
Toraut. A female (22.1.1985) and a male
(6.111.1985) taken beside S. Toraut.
Zyxomma obtusum Albarda (42)
Toraut. Quite plentiful at kitchen pond and one
seen at riverine pond at forest edge, but difficult to
catch (only one male taken). It appeared about
18.00 h at kitchen pond where males established
territories along lengths (about 10 m) of pond
margin which they patrolled by flying back and
forth just a few centimetres above the water sur-
face. Their white colouration rendered them con-
spicuous in the fading light.
Tholymis tillarga (Fabricius) (43)
Toraut, Menado. Males common on kitchen
120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
pond and one observed hovering 50 cm above an
ovipositing female. A female came to an electric
light at 21.00 h and another was found just in the
forest resting beneath a palm frond.
Tramea eurybia (Selys)
Ternate. Two males captured 2.iii.1985 at edge of
volcano crater at 1700 m altitude.
T. transmarina (Brauer) subsp. nov. (44)
Toraut. Single male, 27.1.1985, flying in sunshine
at base camp. Several specimens of a Tramea spe-
cies were seen flying above the road at Danau
Mooat.
Pantala flavescens (Fabricius) (45)
Toraut, Menado. Very common about base camp,
hawking in numbers at 2-4 m above the ground. A
male came to electric light about 22.00 h.
Celebothemis delecollei Ris (46)
Toraut. Both sexes found throughout March at
the pool in S. Tumpah by its confluence with first
tributary. Not observed elsewhere.
GEOGRAPHICAL RANGES AND HABITATS OF THE
TORAUT TAXA
Of the 46 taxa collected within about 2 km of the
Project Wallace base camp on S. Toraut, ten are
apparently undescribed, twelve are so far known
Table 1. Geographical distribution of species and sub-
species by families in the Toraut fauna.
a
x
> S
al fs G
5 ÿ +5 as
3 fe} LS
GETS OSE
Zygoptera
Calopterygidae 1 0 0 1
Chlorocyphidae 2(1) 0 0 3
Lestidae 0 1 0 1
Coenagrionidae 2 3 2 7
Protoneuridae 1(2) 0 0 3
Platystictidae (2) 0 0 2
Anisoptera
Aeshnidae 1 1 1 3
Corduliidae l 0 0 1
Libellulidae 4(5) 11 5 25
Totals 12(10) 16 8 46
Table 2. The distribution by habitats of taxa known
only from Sulawesi and of those with a broader distribu-
tion.
>
uw
Ba SE
os 7 fas
ee AE ire}
gede) oa bed
ud oa i)
A € Na (ce)
Endemic & undescribed taxa 16 4 2
More widespread taxa 3 15 6
only from Sulawesi, 16 are found in the Oriental
region (in some cases in other zoogeographical
regions as well but not in the Australasian region)
and eight have ranges that include both the Orien-
tal and Australasian regions. Ranges of the taxa by
families are shown in table 1. Most of the Zygo-
ptera (11 of 17) are so far known only from Sula-
wesi, but the majority of Anisoptera (18 of 29)
have considerably broader ranges. The level of en-
demism in Sulawesi Odonata is high, but the fauna
includes many Libellulidae, the dominant family,
with a broad distribution in south-east Asia. There
are few indications of Australasian elements in the
odonate fauna. Only the single species of Isosticti-
dae, which was not found at Toraut, has a probable
Australasian derivation. The dragonfly fauna of
Sulawesi is therefore predominantly Oriental, a
conclusion reached also by van Tol (1987a), but
knowledge of dragonfly distribution is insuffi-
ciently complete to allow us to indicate narrower
geographical affinities. The existence of a small
Australian component is not at variance with the
suggestion that Sulawesi is a conglomerate of part
of an island arc of Oriental origin forming Sangir,
the northern peninsula (Sulawesi Utara) and we-
stern Sulawesi, and a mass including the eastern
and south-eastern peninsulas originating in the
Australasian region.
Habitats of the Toraut fauna may be classified as
primary (those in the rainforest and major rivers)
or secondary (those in clearly man-modified situ-
ations). The distribution of taxa between these two
habitat classes (table 2) shows that the endemic
taxa predominate in primary habitats whilst spe-
cies having a broad geographical range occur prin-
cipally in secondary habitats. Whilst this is what
would be expected, the data are influenced by the
absence of permanent ponds, suitable for colonisa-
tion by the more widespread Libellulidae, from the
forest area surveyed. Van Tol (1987a) similarly
found the Odonata of primary forest streams to be
nearly all endemic to Sulawesi. He lists (van Tol
1987a, 1987b) the following additional species
from Dumoga-Bone: Celebargiolestes cinctus (Se-
ASKEW ET AL: Odonata from North Sulawesi
lys) (Megapodagrionidae), ?Paragomphus capita-
tus (Martin) (Gomphidae) — found as larva only,
Nannophlebia aglaia Lieftinck (Libellulidae) and
Diplacina torrenticola van Tol which are all ende-
mic, plus Orthetrum pruinosum (Burmeister) (Li-
bellulidae).
ACKNOWLEDGEMENTS
Provision of a scientific investigations grant by
The Royal Society enabling Askew to participate in
Project Wallace is gratefully acknowledged. This
paper is based in part on material collected on
Project Wallace, sponsored by the Royal Entomo-
logical Society of London and the Indonesian Ins-
titute of Sciences (Results of Project Wallace No.
76). Colin Johnson (Manchester Museum), Steve
Judd and Dr Ian Wallace (Liverpool Museum; Na-
tional Museums and Galleries on Merseyside) allo-
wed Cleland access to collections and use of facili-
ties.
REFERENCES
Kitching, R. L., 1986. A dendrolimnetic dragonfly (Anis-
optera: Libellulidae) from Sulawesi. — Odonatologica
15: 203—209.
Lieftinck, M. A. 1936. A new genus and species of Libel-
lulidae from Celebes (Odon.). — Treubia 15:
399—403.
Van Tol, J., 1987a. The Odonata of Sulawesi (Celebes),
Indonesia: an introduction. — Advances in Odonato-
logy 3: 147—155.
Van Tol, J., 1987b. The Odonata of Sulawesi and adjacent
islands. Parts 1 and 2. — Zoologische Mededelingen,
Leiden 61: 155-176.
Received: 17 October 1988
Accepted: 29 November 1989
121
122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
ANNOUNCEMENT
Fauna Malesiana Foundation
Fauna Malesiana: surveying one of the world's
richest faunas
Zoologists working on the fauna of the Indo-Aus-
tralian archipelago have long felt the need for a
coordinating research body, similar to the Flora
Malesiana Project in botany. Both the archipelago
and the adjacent continental regions possess a lar-
gely unexplored fauna of immense diversity, which
is of great scientific and economic interest to the
nations in the region and indeed the entire world.
Taxonomists, biogeographers and geologists have
lately produced many new, exciting ideas about the
evolution of life in the archipelago. Taxonomists
are also pressing ahead with in-depth studies of
animal and plant groups, frequently supported by
field work, nowadays facilitated by the enhanced
accessibility of poorly explored regions. Since 1980,
terrestrial and marine expeditions have contrib-
uted to the steep increase in our knowledge of
lesser known animal groups inhabiting the ar-
chipelago. Clearly, an increased understanding of
the composition of the fauna and flora will assist
resident policy makers and applied scientists in the
rational management of natural and man-made
ecosystems. The local scientific community, as well
as agriculture, forestry, fisheries and the general
public will no doubt profit from forthcoming re-
views of our present knowledge of the fauna. It is
proposed to publish such reviews under the com-
prehensive title Fauna Malesiana.
Fauna Malesiana Foundation: starting a new or-
ganization
To coordinate and expand the continuing survey of
the fauna of the Indo-Australian archipelago a
foundation has recently been established, the
Fauna Malesiana Foundation, based in Leiden, The
Netherlands. This new organization will be ad-
vised by scientists from various countries, includ-
ing those situated in the region. The Foundation is
currently seeking financial support for several pro-
jects, one being the establishment of a Fauna Male-
siana Bureau. The Bureau will produce a Fauna
Malesiana Bulletin (in a format similar to the Flora
Malesiana Bulletin) and serve as the administrative
centre for the Foundation’s various activities.
Another important project is the production of a
new series of publications, the Fauna Malesiana.
This series is envisaged to comprise handbooks for
the identification of animal groups as well as other
documents relating to the fauna of the archipelago.
The permanent address of the Foundation is:
Fauna Malesiana Foundation, PO.Box 9517,
2300 RA Leiden, The Netherlands.
THE SULAWESI GENUS BRACHYLOBOPYGA (HOMOPTERA:
CICADIDAE)
by
J. P. DUFFELS
ABSTRACT
Duffels, J. P., 1989. The Sulawesi genus Brachylobopyga (Homoptera: Cicadidae). — Tijd-
schrift voor Entomologie 132 (1): 123—127, figs. 1-8. [ISSN 0040-7496]. Published 3 July 1989.
The discovery of the new cicada species Brachylobopyga montana and the alleged synonymy
of B. decorata Duffels, the type species of the genus, to Cicada toradja Breddin prompted a
reconsideration of the genus Brachylobopyga. The distribution of this genus is compared with
the pattern of endemism displayed by its supposed sister genus Dilobopyga.
Key words. — Cicadidae; Brachylobopyga; key; Indonesia; biogeography.
Dr. J. P. Duffels, Instituut voor Taxonomische Zoölogie (Zoölogisch Museum), Universiteit
van Amsterdam, The Netherlands.
INTRODUCTION
The genus Brachylobopyga was erected for a new
species, B. decorata, from Sulawesi (Duffels 1982).
This species was described from Bua Kraeng (1650
m), a mountain peak NW of Gunung Lompobat-
tang in an isolated massif in the southwestern arm
of Sulawesi. Another species of this genus, already
mentioned by Duffels (1986), was recently col-
lected at high altitudes (1600-1900 m) in Central
Sulawesi. This new species is described here as
Brachylobopyga montana.
In connection with my current studies of Su-
lawesi cicadas, I have scrutinized again the species
with uncertain systematic position described by
Breddin (1901). This led to the conclusion that
Brachylobopyga decorata is identical to Cicada to-
radja Breddin, 1901.
TAXONOMY
Brachylobopyga Duffels
Brachylobopyga Duffels, 1982: 156; Duffels 1986:
320—322.
Type species: Brachylobopyga decorata Duffels, 1982 (=
Brachylobopyga toradja (Breddin, 1901)).
The genus was characterized (Duffels 1982) by
a long, strongly chitinized, apically flattened aedea-
gus and a narrow pygofer with parallel lateral sides
and, apically truncate, lateral processes. B. montana
is attributed to Brachylobopyga because of the very
123
similar structure of the aedeagus, though the py-
gofer of this species is widest across the middle (fig.
2) and has short, apically rounded, lateral processes
(fig. 3). B. montana also differs from B. toradja in
the relatively shorter basal pygofer lobes and in
colouration and black body marking. The male
opercula are very alike, viz., very long and narrow-
ing to the apices.
The discovery of the new species of Brachy-
lobopyga shows that the long, strongly chitinized,
apically flattened aedeagus remains a synapo-
morphy for Brachylobopyga, but the short and
rounded lateral process of the pygofer makes less
certain that Brachylobopyga is the sister genus of
Dilobopyga (cf. cladogram of the subtribe Cosmo-
psaltriaria in Duffels 1986: fig. 2).
KEY TO THE SPECIES OF BRACHYLOBOPYGA
Underside of postclypeus with small, median,
black-brown spot at one third from anterior
margin. Ground colour of male abdomen and
timbal covering yellowish brown. Pygofer with
truncate lateral lobes, long basal lobes and pa-
rallel lateral sides (Duffels 1982: figs. 4 & 5).
Clasper long, reaching to halfway the length of
the pygofer, and narrowing to its apex (Duffels,
1982: figs. 4 & 7). Body length @: 25.7—30.2
124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
— Underside of postclypeus with fairly broad, me-
dian fascia. Ground colour of male abdomen
castaneous brown; timbal covering black-
brown. Pygofer with rounded lateral lobes,
short basal lobes and widest across the middle
(figs. 2 & 3). Clasper short, shaped as in fig. 4.
Body length 8: 30—34.5 mm........ montana
Brachylobopyga toradja (Breddin, 1901)
comb. n.
Cicada toradja Breddin, 1901: 109—111.
Cosmopsaltria toradja, Sharp 1903: 279.
Cicada (?) toradja, Distant 1906: 41; Distant 1912: 31.
Tibicen toradja, Kato 1956: 80; Duffels & van der Laan
1985: 78.
Tibicen toradjus, Metcalf 1963: 320.
Brachylobopyga decorata Duffels, 1982: 156—159, figs.
1—8 (syn. n.).
Synonymy. — Breddin described Cicada toradja
from South Celebes (= Sulawesi Selatan): “Wawo-
karaeng, 5000 Fuss, Febr. (Fruhst.)”. The descrip-
tion was probably made from one male specimen
only, since measurements are given for one speci-
men. The type should be in the “Institut fiir Pflan-
zenschutzforschung der Akademie der Landwirt-
schaftswissenschaften der DDR” in Eberswalde,
like the types of all cicadas described in Breddin’s
1901 publication. Unfortunately, the type could not
be located in the Eberswalde collection (pers.
comm. Dr. G. Petersen, 1.vii.1985). This was not
unexpected since the type was not listed in Gaedic-
ke's catalogue of type specimens in the collection
of this institute (1971). Earlier study of two other
cicada types of Breddin revealed that the original
identification labels of Breddin were removed
while new labels with identifications of A. Jacobi
were added (Duffels 1977: 144, 147). This may
have happened also to the type of Cicada toradja.
Breddin’s description of this species perfectly fits
Brachylobopyga decorata especially in the marking
of the body and the characteristic shape of opercu-
lum and pygofer lobes. Locality, altitude, month of
collecting and collector of Breddin’s type of C. to-
radja are identical to those of the type-series of B.
decorata, of which all specimens have identical
labels. It is obvious that these specimens and Bred-
din's specimen of C. toradja are part of the same
series. Breddin (1901: 3) mentioned that he studied
a part [sic !] of the Hemiptera collections made by
H. Fruhstorfer in Minahassa and South Celebes in
1895 — 1896. The considerations given so far lead
to the conclusion that Brachylobopyga decorata is
a junior synonym of Cicada toradja.
Description. — For an extensive description of
this species the reader is referred to the original
descriptions of C. toradja and B. decorata.
Distribution. — Bua Kraeng (= Wawokaraeng),
a mountain peak in an isolated massif in South-
West Sulawesi, SE of Ujung Pandang.
Brachylobopyga montana sp. n.
(figs. 1—8)
Description. Ground colour of head and thorax
light ochraceous, in some specimens with a green-
ish tinge. Ground colour of abdomen brownish.
Head. — Male: Supra-antennal plates for the
greater part black. Vertex with large, round, black
spot enclosing ocelli and reaching from fronto-
clypeal suture to, or nearly to, anterior pronotum
margin, a pair of fairly large triangular spots next
to median spot and a pair of small, oval spots
medio-proximally of eyes. Anterior part of postcly-
peus black, this black part encloses a round, pale
ochraceous spot at anterior postclypeus margin and
continues on underside of postclypeus in a black
median fascia, which is anteriorly as wide as ante-
rior ochraceous spot and narrows strongly to cly-
peal suture. Anteclypeus with, sometimes faint,
brown, median spots at mid-length. Medial margin
of mandibular plate black along anteclypeus. Ro-
strum ochraceous, its dark brown apex just re-
aching trochanter of hind legs.
Female: As in male but spots next to median
spot considerably smaller.
Pronotum. — Male: Markings black. Central fas-
ciae either very narrow or extremely thin or lacking
in the middle, anteriorly strongly widened into
black line along pronotum margin, posterior ends
semicircular. Black lines lie above proximal ends of
anterior oblique fissures. Two pairs of black marks
between both pairs of oblique fissures are broadly
connected with black anterior oblique fissures. Po-
sterior oblique fissures and lateral part of ambient
fissure filled with broad, irregular, black fasciae.
Pronotum collar with weakly developed, very ob-
tuse, latero-distal tooth and a very indistinct, brow-
nish spot on latero-proximal corner.
Female: As in male but spots between oblique
fissures smaller and black colouration of fissures
much narrower.
Mesonotum. — Male: Fasciae black. Median fas-
cia narrow anteriorly, widening to 4—5 times its
anterior width at 3/5 its length and narrowing
again towards cruciform elevation. Paramedian
fasciae converge slightly from anterior mesonotum
margin to half the length of mesonotum disk; they
are anteriorly as broad as anterior part of median
fascia but widen considerably toward their poste-
rior ends. A pair of fairly large spots is situated in
front of anterior angles of cruciform elevation.
DUFFELS:
Areas between median and paramedian fasciae
chocolate brown. Lateral fasciae broad with some
irregular black spots next to its anterior part; fas-
ciae embedded in chocolate brown. Cruciform ele-
vation yellow ochraceous, apices of its anterior
arms black.
Female: Marking less developed. Anterior half
of lateral fasciae clouded with brown.
Legs. — Light brown. Fore femur with a black
line along its underridge, which connects a proxi-
mal, fairly long, appressed, dark spine and a shor-
ter, more distally placed, erect, dark spine. Upper-
sides and innersides of femora with longitudinal,
brown lines. Underside of fore tibia and its distal
end all around dark brown; middle tibia dark brown
apically. Tarsi of fore and middle legs blackish
brown.
Tegmina and wings. — Hyaline. Venation of
tegmen light brown variegated with brown-black
parts, and turning brown-black apically. Venation
of wing dark brown. Tegmina with fairly broad
infuscations at bases of 2nd and 3rd apical areas;
two paratypes also very lightly infuscated at apices
of veins of 2nd, 3rd (and áth) apical areas.
Operculum. — Male (fig. 7): Castaneous and
long, reaching posterior margin of sixth abdominal
segment. Surface of operculum fairly convex and
rugose. Operculum at its widest close to its base and
tapering to fairly narrow and rounded apex. Medial
margin nearly straight but very slightly concave at
two thirds of its length in holotype and one para-
type. Lateral margin weakly convex.
Female (fig. 8): Light brown, just reaching po-
sterior margin of abdominal segment 2. Latero-
proximal corner black. Lateral margin basally sinu-
ate. Latero-distal angle rounded. Posterior margin
almost straight, but slightly curved toward mera-
canthus.
Abdomen. — Male: Ground colour castaneous
brown. Timbal coverings blackish brown (fig. 6).
Segment 2 with more or less distinct, blackish
brown, middorsal triangle at anterior margin. An-
terior and posterior margins of segments 2 and 7
dorsally dark brown. Segments 3—6 with a pair of
sublateral dark spots at half the segment’s height.
Distal half of segment 7 ochraceous. Ventral side
of abdomen castaneous, posterior margins of seg-
ments 3—5 somewhat darker brown.
Female: Brownish with scattered pilosity on dor-
sal surface. Posterior margin of segment 2 black.
Segments 2—4 with black transverse marks at po-
sterior margins and similar but smaller marks on
segments 5—7. Segments 4—7 with sublateral,
round black spots. Segment 9 with two paramedian
obconical black marks.
Brachylobopyga 125
Fig. 1. Brachylobopyga montana, holotype.
Male genitalia. — Lateral processes of pygofer
short and apically rounded (fig. 3). Cup of pygofer
formed by the basal lobes U-shaped; basal pygofer
lobes distinctly shorter than in B. toradja (fig. 2).
Clasper with apical and subapical hooks, both di-
rected laterad (fig. 4). Aedeagus chitinized, very
long and strongly curved, apex flattened and poin-
ted (figs. 3 & 5).
Measurements. — Body length & 30—34.5 mm,
Q 25.0 mm; head width & 9.5—10.0 mm, Q 9.1
mm; pronotum width ¢ 9.7—10.7 mm, 2 9.6 mm;
mesonotum width ¢ 8.7—9.2 mm, 9 8.5 mm; teg-
men length & 40.6—42.0 mm, 9 40.0 mm.
Holotype, 8, “Indonesia, Sulawesi Tengah / Lore
Lindu N.P.”, “10 km SE Poloka / 1900 m, 25.iii.1985 / J.
P.& M.J. Duffels”, “Stat. 55 / Disturbed lower / montane
forest ML-light, canopy’. Paratypes, Indonesia, Sulawesi
126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 2—8. Brachylobopyga montana. — 2, pygofer in ventral view, holotype; 3, apical part of pygofer in lateral view,
holotype; 4, right clasper in lateral view, holotype; 5, apical part of aedeagus, paratype Puncak Dingin; 6, left timbal
covering, paratype Puncak Dingin; 7, male abdomen with operculum in ventro-lateral view, holotype; 8, female
operculum in ventro-lateral view, paratype.
Tengah: 1 &, same data as holotype; 1 6, Lore Lindu N.P.,
Rano Rano, 1600 m, 10 km NE of Gimpu, 15.iii.1985, J.
P. & M. J. Duffels, stat. 43, Lower montane forest, ML-
light; Puncak Dingin, ca. 1700 m, 15.x.1985, S. Nagai leg.,
1 9, same data but 21.x.1985, 1 @.
Other material (freshly emerged, not fully coloured
and somewhat crushed): Puncak Dingin, ca. 1700 m,
15.x.1985, S. Nagai leg., 1 @, same data but 21.x.1985, 1
8, 15.xi.1985, 1 &. The holotype and the paratype from
Rano Rano are deposited in the Instituut voor Taxonomi-
sche Zoölogie (Zoölogisch Museum), Amsterdam; the
other paratype from Poloka is in the Museum Zoologi-
cum Bogoriense, Bogor; the material from Puncak Din-
gin is in the collection of Dr. M. Hayashi, Urawa, Japan.
Distribution. — The species is recorded from
lower montane forest in Sulawesi Tengah. Rano
DUFFELS:
Rano (1600 m) is situated in the mountain range
east of the valley of Salo Palu, 85 km South of Palu
and 10 km E of Gimpu. Poloka (1900 m) is situated
along the road from Palu to Sedeo, about 65 km SE
of Palu. This is very close to, or perhaps about the
same locality as, Puncak Dingin (ca. 1700 m),
which is on the southern slope of Mt. Roreka-
ofimbu (M. Hayashi, pers. comm.).
BIOGEOGRAPHY
The supposed sister genus of Brachylobopyga is
Dilobopyga, which is distributed in Sulawesi inclu-
ding Muna and Buton (with 30 species), Sangihe Is.
(1 species), Selayar (1), Banggai archipelago (2),
Sula Is. (1) and South Maluku (1) (Duffels 1986, in
press). Up to now, only ten out of these 36 species
of Dilobopyga have been described. Species of
Dilobopyga are found from sea-level to 1600 m
mainly in rainforest habitats, though some species
are found in other vegetation types and gardens.
Preliminary phylogenetic studies of this genus
show that monophyletic groups are centred in
North and Central Sulawesi, inSW Sulawesi and in
East Sulawesi.
The two species of Brachylobopyga are found in
high altitude rainforest areas (1600—1900 m). B.
toradja seems to be restricted to the isolated moun-
tain massif of SW Sulawesi. From this area only
one endemic Dilobopyga species (D. margarethae
Duffels) and one widespread species (D. minahas-
sae (Distant)) have been recorded. B. montana is
found in Central Sulawesi, where several species of
Dilobopyga and one or two monophyletic species
groups of this genus have a very restricted distri-
bution.
ACKNOWLEDGEMENTS
I am very much indebted to Ir. Syafii Manan
(Directorate General of Forest Protection and Na-
ture Conservation, Bogor) for permission to collect
in Lore Lindu National Park, Sulawesi Tengah. lam
grateful to Mr. Rolex Lameauda and Mr. Julian
Sango (Lore Lindu National Park), to Mr. Frans
Mamarimbing and Mr. Boy Manopo (Hotel Vis-
cana, Palu), to our porters Jonathan, Kantoro and
Obed from the village of Gimpu and to my wife
Greet Duffels-van Egmond for their help in the
field. Dr. M. Hayashi (Saitama University, Urawa,
Japan) kindly sent me his material collected by Mr.S.
Nagai. I thank Mr. A. J. de Boer for criticizing the
manuscript of this paper, Mr. D. Langerak for the
preparation of fig. 1 and Mrs. Annelies Stoel for
typing the manuscript. The fieldwork was funded in
part by the Netherlands Foundation for the Advan-
cement of Tropical Research (WR 85—197).
Brachylobopyga 127
REFERENCES
Breddin, G., 1901. Die Hemipteren von Celebes. Ein
Beitrag zur Faunistik der Insel. — Abhandlungen der
Naturforschenden Gesellschaft zu Halle 24: 1—213,
pl 1.
Distant, W. L., 1906. A synonymic catalogue of Homo-
ptera. Part 1. Cicadidae. Trustees of the British Mu-
seum, London.
Distant, W. L., 1912. Homoptera, Fam. Cicadidae, Sub-
fam. Cicadinae. — Genera Insectorum 142: 1—64, pls.
1—7.
Duffels, J. P., 1977. A revision of the genus Diceropyga
Stal (Homoptera, Cicadidae). — Monografieén van de
Nederlandse Entomologische Vereniging 8: 1—227.
Duffels, J. P., 1982. Brachylobopyga decorata n. gen., n. sp.
from Sulawesi, a new taxon of the subtribe Cosmo-
psaltriaria (Homoptera, Cicadoidea: Cicadidae). —
Entomologische Berichten, Amsterdam 42:
156—160.
Duffels, J. P., 1986. Biogeography of Indo-Pacific Cicado-
idea: a tentative recognition of areas of endemism. —
Cladistics 2: 318—336.
Duffels, J. P., in press. Biogography of Sulawesi cicadas
(Homoptera: Cicadoidea). — Proceedings Sympo-
sium Royal entomological Society.
Duffels, J. P. & Laan, P. A. van der, 1985. Catalogue of the
Cicadoidea (Homoptera, Auchenorhyncha)
1956-1980. — Series Entomologica 34; i—xvi,
1—414.
Gaedicke, H., 1971. Katalog der in den Sammlungen des
ehemaligen Deutschen Entomologischen Institutes
aufbewahrten Typen-VI. — Beiträge zur Entomologie
21:35 339:
Kato, M., 1956. The biology of cicadas. Bulletin Cicadidae
Museum. Iwasaki Shoten, Jinbocho Kanda, Tokyo.
Metcalf, Z. P., 1963. General catalogue of the Homoptera
8. Part I Cicadidae, Section I Tibiceninae. North Ca-
rolina State College, Raleigh, N.C.
Sharp, D., 1903. XIII Insecta. — The Zoological Record
(1902) 39: 1—313.
Received: 20 February 1989
Accepted: 3 March 1989
128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
BOOK ANNOUNCEMENTS AND REVIEWS
Holmen, M., 1987. The aquatic Adephaga (Coleoptera) of
Fennoscandia and Denmark, I. Gyrinidae, Haliplidae,
Hygrobiidae and Noteridae. — Fauna Entomologica
Scandinavica 20: 1-168, 288 figs, 1 map. [ISBN 90 04
08185 2, E. J. Brill, Leiden, the Netherlands. Price 64
Dutch Guilders/US $ 32.— |.
This volume treats the northern European re-
presentatives of the small aquatic beetle families
mentioned in the title, which together with the
large Dytiscidae (the subject of a future volume)
and the non-European Amphizoidae, form the
aquatic Adephaga. Apart from the 36 species occur-
ring in Northern Europe, the volume treats 5 ad-
ditional species, so covering most of West and
Central Europe. Apart from extensive keys, the
book provides concise descriptions, with informa-
tion on distribution, biology and nomenclature.
Some general information on morphology, zoo-
geography and biology is provided with each fam-
ily. One colour plate illustrates the general aspect
of the genera treated. As in other volumes of the
series, the distribution over the provinces of the
four northern countries, is listed in a dot catalogue.
This book will be highly welcomed by workers
on aquatic beetles and aquatic ecologists, since no
recent reliable keys to these small, but difficult
families were available. Where previous keys al-
most all suffer from poor illustrations and absence
of detailed information on genitalia, this book is
very well illustrated with the use of a wealth of
diagnostic characters, including male and female
genitalia and stereoscan pictures of elytral sculp-
ture. The information on biology is also much more
extensive than one usually finds in identification
guides, and includes amongst others information
on larva and pupa. The author also indicates where
the description of the larva of each species can be
found.
One minor point of criticism is that the author,
while fixing lectotypes for various species, failes to
solve the nomenclatorial confusion around
Haliplus lineolatus Mannerheim, by designating a
lectotype, although he points out himself that this
ought to be done.
Mr Holmens publication is highly recom-
mended, and we can hardly wait to see the volume
on Dytiscidae.
[E. J. van Nieukerken]
Duffels, J. P., 1988. The Cicadas of the Fiji, Samoa and
Tonga Islands, their Taxonomy and Biogeography
(Homoptera, Cicadoidea) with a chapter on the geo-
logical history of the area by A. Ewart. — Entomono-
graph 10: 1—108, figs. 1—207 [E. J. Brill, Leiden, The
Netherlands. Price 56 Dutch Guilders = c.
US$ 28.00].
With this book the author continues his studies
on the cicadas of the oriental and australian region.
The adults of 24 (sub) species occurring in the Fiji,
Rotuma, Samoa and Tonga Islands are thoroughly
described and illustrated. Also descriptions of lar-
vae and exuviae, as far as known, and details on
behaviour and distribution are provided. The phy-
logenetic relationship of the species is extensively
discussed also in relation to other taxa of the
southwest Pacific. Biogeographic patterns are an-
alysed and compared with the geological history of
the area, which is described in a chapter by A.
Ewart.
This publication maintains the high standard as
was set by the author in his previous publications.
The descriptions are concise but fully adequate. I
also appreciate the illustrations. Habitats and liv-
ing insects are photographed, while diagnostic
characters are illustrated with line drawings. The
quality of the figures of the whole insects (without
their wings), prepared by J. Zaagman, is particu-
larly high.
This book is, of course, a must for the specialists
in this group of conspicuous Homoptera, but its
value for other zoologists working in the Pacific on
biogeography should also be stressed. It is a most
stimulating addition to our knowledge of this area.
[J. van Tol]
DIE GATTUNG POLEMIOSILIS PIC AUF SULAWESI
(COLEOPTERA, CANTHARIDAE)
(41. Beitrag zur Kenntnis der indo-malaiischen Cantharidae)
von
W. WITTMER
ABSTRACT
Wittmer, W., 1989. Die Gattung Poleomiosilis Pic auf Sulawesi (Coleoptera, Cantharidae).
(41. Beitrag zur Kenntnis der indo-malaiischen Cantharidae). — Tijdschrift voor Entomologie
132: 129-134, figs. 1-13. [ISSN 0040-7496]. Published 3 July 1989.
The species of Polemiosilis Pic occurring on Sulawesi are revised. Six species and one
subspecies are listed of which five species are described as new to science. Polemiosilis tes-
taceobasalis Pic is transferred to Gwineapolemius and Gwineapolemius tolianus (Pic) is syn-
onymized with P. testaceobasalis Pic.
Key words. — Cantharidae, Sulawesi.
Dr. W. Wittmer, Naturhistorisches Museum, CH-4001 Basel, Schweiz.
DIE GATTUNG POLEMIOSILIS PIC
Von dieser Gattung sind bisher zwei Arten von
Sulawesi beschrieben worden: P. rufosuta Pic, 1921
and P. testaceobasalis Pic, 1921, sowie P. saleie-
rensis Wittmer, 1973 von der Insel Saleier, die
zum Faunengebiet von Sulawesi gehört. Die Art
P. testaceobasalis Pic is zu streichen, sie wird in
die Gattung Guineapolemius versetzt, siehe letzten
Abschnitt.
Das reiche zur Verfügung stehende Material
verdanke ich Herrn Dr. Peter Hammond, London;
es stammt aus den Aufsammlungen des "Project
Wallace” (unterstützt durch die Royal Entomo-
logical Society of London und die Lembaga Ilmu
Pengetahuan Indonesia), und dasjenige welches
mir von Herrn Prof. Dr. Masataka Satö, Nagoya,
aus den Aufsammlungen von Herrn Shinji Nagai
zugestellt wurde. Beiden Herren danke isch be-
stens für die Zustellung des Materials. Des wei-
teren danke ich Herrn G. Hodebert, Paris, für die
sorgfältig ausgeführten Zeichnungen.
Verwendete Abkürzungen: BM = British Mu-
seum (Natural History), London; MP = Museum
National d’Histoire Naturel, Paris; NHMB = Na-
turhistorisches Museum, Basel.
Results of Project Wallace, no. 77.
SCHLÜSSEL
Die auf Sulawesi vorkommenden Arten der Gat-
tung Polemiosilis & lassen sich wie folgt unter-
scheiden:
1. Flügeldecken zum grössten Teil schwarz, höch-
stens an der Naht ein wenig aufgehellt.... 2
Flügeldecken zum grösten Teil gelbbraun, Ba-
sis verschwommen angedunkelt, Spitzen nicht
sehr breit schwarz, manchmal ist die basale
Verdunkelung längs der Seiten schmal mit den
Spitzen verbunden, oder die Basis der Flügel-
decken ist nicht angedunkelt und lediglich die
Seiten unter den Schultern sind schwach dun-
kein P. rufosuta ssp. saleierensis Wittmer
. Hälfte des letzten Sternits mit 1 oder 2 Dornen
oder Verlängerungen. Halsschild einfarbig
orange
Hälfte des letzten Sternits ohne Dornen oder
Verlängerungen. Halsschild dunkelbraun, nur
der Basal- und Vorderrand schmal oder die
basale Hälfte aufgehellt, selten ist der Hals-
schild einfarbig orange. Abb. 2
re u ona P. simplex n.sp.
. Hälfte des letzten Sternits mit 2 Dornen oder
Merlanserungene ep re RER
Hälfte des letzten Sternits mit nur 1 Dorn,
130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Abb. 1, 3, 5: Kopulationsapparat halb seitlich von Polemiosilis &: 1, simplex n.sp. 3, rubidiceps n.sp. 5, hammondi
n.sp. — Abb. 2, 4, 6: Hälfte des letzten Sternits von Polemiosilis &: 2, simplex n.sp. 4, rubidiceps n.sp. 6, hammondi
n.sp. Massstab für Abb. 1-6.
Kopf einfarbig orange, nur neben den Augen
mehr oder weniger angedunkelt. Abb. 4 .....
RBE AAT AE P. rubidiceps n.sp.
4. Hälfte des letzten Sternits mit 2 ziem-
lich regelmässig geformten Dornen verse-
Lene: FUN TENTEN PY TEUGE Ri, 5
— Hälfte des letzten Sternits mit 1 Dorn und
einer Verlingerung versehen, die auch spitz
Sciare OR ele 6
5. Die beiden Dorne sind sehr kurz, der obere
überragt den äusseren Rand des Seitenteils
nichtodenkaum AbhAC RENE AA a
> RA eit pe tonnara) P. hammondi n.sp.
— Die beiden Dorne sind sehr lang, auch der
obere überragt den aüsseren Rand des Seiten-
taillé, Alan, OÙ ss re EU P rufosuta Pic
6. Der obere Dorn entspringt am Hinterrand des
Sternits und ist hakenartig gekriimmt, die Ver-
längerung darunter ist sehr breit und bildet
einenkspitzenmWinkel Ap blj
TRAE, SR NEO ENE SAIS P. nagati n.sp.
— Der obere Dorn entspringt neben der Mitte, er
ist nur leicht gekriimmt, die Verlängerung da-
runter ist breiter als der Dorn, Spitze gerundet.
Abba la. ee P. nanggalaensis n.sp.
Polemiosilis simplex sp.n.
(Abb. 1-2)
Holotypus: Indonesia, Sulawesi Utara: Danau Mooat
near Kotamobagu, 1200 m, 16.-19.11.1985 und xi.1985,
plants by lake. (BM).
Paratypen: als Holotypus, 1 BM, 2 NHMB; Tondano
(Celebes), 1 4 NHMB.
Kopf schwarz, vor den Fiihlerwurzeln aufge-
hellt, selten ganz schwarz; Fühler schwarz, oder die
beiden Glieder und die Spitze des letzten Gliedes ist
leicht aufgehellt; Halsschild dunkelbraun, nur der
Basal- und Vorderrand schmal, oder die ganze ba-
sale Hälfte ist aufgehellt, selten ist der ganze Hals-
schild orange; Schildchen orange bis dunkelbraun;
Flügeldecken schwarz; Beine dunkel, bei dem Ex-
WITTMER:
emplar von Tondano mit orangenem Halsschild
sind alle Schenkel und Schienen orange, nur die
Knie leicht angedunkelt.
8. Kopf mit den Augen breiter als der Hals-
schild, Stirne leicht gewölbt, Basis bis zur Mitte der
Augen deutlich und dichter punktiert als weiter
vorne. Fühler ungefähr so lang wie die Flügeldec-
ken, von Glied 3 an platt, 3 bis 10 gegen die Spitze
verbreitert. Halsschild breiter als lang (47X38),
Seiten kurz vor der Mitte am breitesten, hier ge-
rundet, dann gegen die Basis schwach verengt, Ba-
salecken ein wenig vorstehend, schwach stumpf-
winklig; Scheibe mit 2 schwachen Beulen, die durch
eine feine Längslinie voneinander getrennt sind,
die besonders an der Basis gut sichtbar ist, Oberflä-
che fast glatt, feine Haarpunkte sichtbar (64X).
Flügeldecken parallel, körnig gewirkt, mit Spuren
von | bis 2 Langsrippen, zerstreute Punkte sicht-
bar. Hälfte des letzten Sternits Abb. 2. Dasselbe
fällt durch seine einfache Form auf, da Dorne oder
Fortsätze vollständig fehlen.
Kopulationsapparat Abb. 1 fast im Profil. Die
Seite des dorsalen Basalstücks ist leicht gerundet
vorgezogen, die Basophysen sind kürzer als das-
selbe, deren Spitze ist fast eckig, der Zahn darunter
liegt verhältnismässig weit vorne.
Länge: 7-8 mm.
Durch die einfache Form des letzten Sternits
sehr charakterisierte Art. Die dunkeln Exemplare
erinnern in der Färbung an P. obscuricolor Pic von
Borneo.
Polemiosilis rubidiceps sp.n.
(Abb. 3-4)
Holotypus: Indonesia, Sulawesi Utara: Dumoga Bone
N.P., Hog’s Back’ Camp, lowland forest, 492 m, xi.-
x11.1985 (BM).
Paratypen: Als Holotypus, 10 BM, 9 NHMB; und:
“Clarke” Camp, lower montane forest, 1140 m, tit
v.1985, 6 BM, 4 NHMB; idem "Edward’s Camp”, lowland
forest, 664 m, x.1985, 6 BM, 6 NHMB; idem site 1, 200
m, New Base Camp site, 27.-28.1.1985, J. D. Holloway, 1
BM; site 2, 200 m, Toraut Bank, 29.-31.1.1985, J. D. Hol-
loway, 1 BM, 1 NHMB; sites 10 und 11, 664 m, Tumpah
Transect, 19.-25.ii.1985, J. D. Holloway, 4 BM, 3 NHMB;
idem G. Mogogonipa summit, 1008 m, v.1985, 1 BM, 1
NHMB, (all the material from Project Wallace expedi-
tion); Sampraga, 28.x.-5.xi.1985 und 17.vi.1986, S. Nagai,
3 coll. Masataka Satò, 3 NHMB.
Kopf rotorange, nur um die Augen schmal ge-
schwärzt, Maxillarpalpen dunkel, oder nur die
Spitze bis das ganze letzte Glied aufgehellt; Fühler
schwarz, Glieder 1 und 2 ein wenig aufgehellt,
letzte 2 bis 3 weisslich, manchmal auch die Spitze
Polemiosilis auf Sulawesi 131
des viertletzten; Halsschild, Schildchen und Beine
orange, alle Tarsen dunkel, meistens auch alle Knie
und oft die Tibien mehr oder weniger angedunkelt;
Flügeldecken schwarz.
8. Kopf mit den Augen breiter als der Hals-
schild, an der Basis mit deutlichen Haarpunkten,
weiter vorne fast glatt. Fühier kräftig, Glieder von
3 an platt, 3 bis 10 gegen die Spitze verbreitert.
Halsschild breiter als lang, Seiten gegen die Basis
schwach verengt; Scheibe mit 2 undeutlichen Beu-
len, die manchmal durch einen Längseindruck ge-
trennt sind, Oberfläche glatt, Haarpunkte zer-
streut, sehr fein punktiert. Flügeldecken praktisch
parallel, körnig gewirkt, mit Spuren von 1 bis 2
Längsrippen. Hälfte des letzten Sternits Abb. 4.
Kopulationsapparat Abb. 3 fast im Profil. Die
Seite des dorsalen Basalstiicks ist in einen fast drei-
eckigen Fortsatz ausgezogen, die Basophysen über-
ragen dasselbe ein wenig.
Länge: 8-9.5 mm.
Neben P rufosuta Pic zu stellen.
Polemiosilis hammondi sp.n.
(Abb. 5-6)
Holotypus: Indonesia, Sulawesi Utara: Dumoga Bone
N.P, “Hog's Back” Camp, lowland forest, 492 m, xi.-
x11.1985, (BM).
Paratypen: als Holotypus, 7 BM, 8 NHMB; und: idem
“Edward’s Camp”, lowland forest, 664 m, iv.1985, 1 BM,
| NHMB; “Clarke” Camp, lower montane forest, 1140 m,
iii. und v.1985, 1 BM, 1 NHMB; G. Mogogonipa, summit,
1008 m, v. und xi.1985, 1 BM, 1 NHMB; site 7 und 8,
Tumpah Transect, 540 m, 9.-11.11.1985 und 12.-13.11.1985,
J. D. Holloway, 1 BM, 1 NHMB (all the material from
Project Wallace expedition).
Firbung wie P. rubidiceps mit Ausnahme des
einfarbig schwarzen Kopfes, der höchstens am
Vorderkopf schwach aufgehellt ist; Fühler eben-
falls dunkler, indem die Glieder 1 und 2 ganz
schwarz sind; manchmal ist die Naht an den Flü-
geldecken schwach aufgehellt.
Sonst wie P. rubidiceps gebaut. Hälfte des letzten
Sternits Abb. 6.
Kopulationsapparat Abb. 5 fast im Profil. Die
Laterophysen erreichen den Rand des dorsalen Ba-
salstiicks nicht, die Seiten dieses sind breit, jedoch
nicht stark aufgebogen. Der seitliche Zahn auf der
Dorsalseite jeder Laterophyse ist fast ganz ver-
deckt.
Lange: 7.5-8.5 mm.
Meinem werten Kollegen Dr. Peter Hammond,
British Museum, London, gewidmet.
Neben P. rufosuta Pic zu stellen.
132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Abb. 7-9: Polemiosilis rufosuta (Pic) 8: 7-8 Kopulationsapparat: 7, von der Ventralseite. 8, half seitlich. 9, Hälfte des
letzten Sternits. Massstab für Abb. 7-9.
Polemiosilis rufosuta Pic
(Abb. 7-9)
Polemiosilis rufosuta Pic, 1921: 26. — Wittmer 1957:
570, Abb. 7.
Der Holotypus befindet sich im MP mit Etikette
“Samanga, S Celebes, xi.1895, H. Fruhstorfer”. Für
die Abb. 9 des letzten Sternits (Hälfte) und des
Kopulationsapparates (Abb. 7-8) diente der Holo-
typus. Ich besitze weitere Exemplare von SW Ce-
lebes: Bonthain, ix.1938, J. P. A. Kalis und Banti-
im NHMB.
Weiterer Fundort: Sulawesi Tengah, near Mora-
wali, Ranu River area, 27.i.-20.iv.1980, M. J. D.
Brendell, 10 BM, 2 NHMB.
Für die Abbildungen 7-9 wurde 1 @ von Bont-
hain verwendet, das mit dem Holotypus von Sa-
manga (MP) übereinstimmt.
Die von mir (Wittmer 1957) veröffentlichte
Abb. 7 des Kopulationsapparates zeigt denselben
von der Dorsalseite und nicht wie angegeben von
der Ventralseite.
P. rufosuta ssp. saleierensis Wittmer stat.n.
Polemiosilis saleierensis Wittmer, 1973, Mitt. Schweiz.
Ent. Ges. 46: 191, Abb. 25.
Nachdem der Holotypus von P. rufosuta Pic auf-
gefunden wurde, und saleierensis hauptsächlich
durch die helle Färbung der Flügeldecken abweicht,
sehe ich mich veranlasst, die Tiere von der Insel
Saleier als Rasse von P. rufosuta Pic zu betrachten.
WITTMER:
Polemiosilis nagaii sp.n.
(Abb. 10-11)
Holotypus: Sulawesi, Peleng I: Luksag, ca. 300 m,
25.ii.-4.iii.1986, Shinji Nagai, in coll. Prof. M. Satô, Na-
goya.
Paratypen: Als Holotypus, in coll. Satô und NHMB.
Kopf einfarbig schwarz, Fühler ebenso, nur die
3 letzten Glieder sind weisslich; Halsschild und
Schildchen orange; Fihler und Beine schwarz, mit
Ausnahme der Schenkel, die an der Basis, manch-
mal bis über die Mitte orange sind.
8. Kopf mit den halbkugelförmigen Augen brei-
ter als der Halsschild, Stirne zwischen den Augen
flach, Oberfläche glatt, zerstreute Haarpunkte vor-
handen. Fühler um fast 1/5 länger als die Flügel-
decken, von Glied 3 an platt, Glieder 3 bis 10 gegen
die Spitze verbreitert, 10 so lang wie 4. Halsschild
breiter als lang (47X39); Seiten knapp vor der
Mitte am breitesten, nach vorne stärker gerundet
als gegen die Basis; Beulen auf der Scheibe nur
schwach entwickelt, ohne Längslinie, die sie trennt,
glatt, Haarpunkte kaum wahrnehmbar. Flügeldec-
ken parallel, fein körnig gewirkt, Längsrippen
kaum angedeutet. Hälfte des letzten Sternits Abb.
11. Der obere Dorn entspringt am Hinterrand des
letzten Sternits, er ist hakenartig nach aussen ge-
kriimmt, die Verlingerung darunter ist sehr breit
Polemiosilis auf Sulawesi 133
und bildet einen kurzen spitzwinkligen Vorsprung.
Kopulationsapparat Abb. 10 fast im Profil. Die
Seite des dorsalen Basalstücks ist in einen fast drei-
eckigen Fortsatz ausgezogen, ähnlich wie bei
rubidiceps und die Basophysen überragen ebenfalls
das dorsale Basalstück.
Q. Fühler ein wenig kürzer, nur ca. 10% länger
als die Flügeldecken, Augen ein wenig kleiner.
Länge: 7-8 mm.
Est freut mich, diese Art ihrem Entdecker Herrn
Shinji Nagai widmen zu dürfen.
Neben P. rubidiceps Wittmer zu stellen.
Polemiosilis nanggalaensis sp.n.
(Abb. 12-13)
Holotypus: S Celebes: Nanggala, Rantepao, 900 m,
v.1938, F. C. Drescher (ex. coll. Wittmer in NHMB).
6. In der Färbung mit P. rufosuta Pic übereins-
timmend, nur daß alle Schienen schwarz sind und
die helle Naht der Flügeldecken fehlt, letztere sind
einfarbig schwarz.
Kopf mit normal grossen Augen, Fühler und
Halsschild wie bei rzbidiceps. Hälfte des letzten
Sternits Abb. 13, dieselbe ist nach aussen in 2 Fort-
sätze ausgezogen, die obere ist schmäler, scharf
spitzig, die untere breiter mit stumpfer Spitze.
Abb. 10, 12: Kopulationsapparat halb seitlich von Polemiosilis &: 10, nagaii n.sp. 12, nanggalaensis n.sp. — Abb. 11,
13: Hälfte des letzten Sternits von Polemiosilis &: 11, nagati n.sp. 13, nanggalaensis n.sp. Massstab für Abb. 10-13.
134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Kopulationsapparat Abb. 12. Das dorsale Bas-
alstiick ist in der Mitte besonders tief gespalten, die
Seiten sind sehr flach, nur wenig erhöht. Die Spit-
zen der Laterophysen erreichen den Apikalrand
des dorsalen Basalstücks nicht.
Länge: 8.5 mm.
Neben P rubidiceps Wittmer zu stellen.
Guineapolemius testaceobasalis (Pic) comb.n.
Polemiosilis testaceobasalis Pic, 1921: 26.
Guineapolemius tolianus Pic, 1921: 27, syn.n.
Bei G. tolianus (Pic) handelt es sich um eine
Farbvariation von G. testaceobasalis (Pic), so dass
tolianus in Synonymie verfällt. Beide Arten sind
von Toli Toli (N Celebes) beschrieben; die Typen
sind im MP.
LITERATUR
Pic, M., 1921. Nouveautés diverses. — Mélanges exotico-
entomologiques 33: 1—32.
Wittmer, W., 1957. 18. Beitrag zur Kenntnis der indo-
malayischen Malacodermata (Col.) (2. Teil). — An-
nals and Magazine of Natural History (12) 10:
567—578, figs. 1—10.
Wittmer, W., 1973. 31. Beitrag zur Kenntnis der indo-
malaiischen Cantharidae (Col.). — Mitteilungen der
Schweizerischen Entomologischen Gesellschaft 46:
175—194, figs. 1—32.
Received 30 March 1989
Accepted 11 April 1989
THE EGG LAYING AND LARVAL DEVELOPMENT OF PHOLCUS
PHALANGIOIDES (FUESSLIN) (ARANEAE: PHOLCIDAE)
by
Ee Gee PLATE
ABSTRACT
Platel, Th. G. H., 1989. The egg laying and larval development of Pholcus phalangioides
(Fuessling) (Araneae: Pholcidae). — Tijdschrift voor Entomologie 132: 135—147, figs. 1-
12, tabs. 1-4. [ISSN 0040-7496]. Published 3 July 1989.
A description is given of the egg-laying of Pholcus phalangioides (Fuesslin). The posture
of the female, when laying eggs, differs from the resting posture. The eggs appear together
with a drop of clear liquid. For a short while the female is kneading this drop after which
it is suspended on a thread and spun. After having done so, she takes the egg-batch in her
chelicerae and retreats to her resting place. The egg-laying behaviour of Ph. phalangioides
is compared with that of Ph. opilionoides (Schrank) and Ph. muralicola Maughan & Fitch.
In captivity the eggs are laid between the end of April and the end of September. A well-
fed female lays her first eggs 10-14 days after the first mating. If she has copulated before,
the moment of egg-laying is determined by the availability of food. Sperm can be stored
by the female. Non-fertilized eggs are removed from the web by the female. The mean number
of eggs is 29.8 (7-63). The maximum number of egg-batches is 9. The mean duration of
the development until the emergence of the first young is 27.2 days. The duration is temperature-
dependent. The female does not hold her eggs continuously in her chelicerae during this
period. Ten to fifteen days before hatching the chorions break and the prelarvae are partially
visible. The female then starts to loosen the threads around the egg mass and helps the
spiderlings to get free. The emerging spiderlings are prenymphs. Their morphology and be-
haviour are described. The prenymphs moult after 7.9 days, dependent on the temperature.
Key words. — Pholcidae; reproduction; behaviour; development.
Th. G. H. Platel, Rozengaard 7, 5283 GD Boxtel, The Netherlands
INTRODUCTION
Since 1981 specimens of Pholcus phalangioides
(Fuesslin) were kept in captivity in order to study
their mating behaviour and other aspects of the
reproduction.
Hardly anything is known in the literature about
egg-laying of this species, although the description
of this phenomenon by Pholcus opilionoides by
Pokrowsky (1899) could refer to Ph. phalangioides.
Maughan (1978) gives a description of the egg
laying with Ph. muralicola (Maughan & Fitch).
As to the development of eggs and young, additions
can be given to the observations published so far.
MATERIAL AND METHODS
The spiders were individually kept in glass cages
(20X15X20 cm) with a wooden lid with an op-
ening that can be closed and a larger one, gauze
covered, for air. The cages were placed in a room
135
on the south-side of my house. The room has par-
tially a flat roof and a large window. Thanks to
this the temperature in the room is strongly de-
pendent on the outdoor temperature. In summer-
time and in sunshine temperatures higher than
30 °C are reached. In cold weather conditions the
room is heated in the daytime but the temperature
is rarely higher than 18 °C and falls back to 10 °C
at night and in case of severe frost even to 5 °C.
The temperatures mentioned in this study were
calculated by means of the monthly survey of the
Koninklijk Nederlands Meteorologisch Instituut
(Royal Dutch Meteorological Service) at De Bilt
(Province of Utrecht), averaging the day-temper-
atures from Beek (Province of Limburg) and De
Bilt.
The spiders were fed with Tipulid midges, Cal-
liphorid and Muscid flies, queens of the ant Lasius
niger about to take to the air and, in winter time,
136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
with maggots or flies just emerged (Calliphoridae).
Unless stated otherwise, all specimens have
been captured at home as adults. Therefore, it was
not known if and how often these females already
laid eggs before. In the cages mainly females were
kept. The males that were put with the female
usually stayed with her until their death. Someti-
mes they were removed after mating.
RESULTS
Egg laying, a case study
The spiders kept in captivity, produced 32 egg-
batches, only two of them were produced in the
daytime. A description of my only complete obser-
vation is given here.
April 21, 1983, 14.45 h. The female hangs upside
down in her web. Some time before she starts
laying eggs, the abdomen is kept horizontally, but
the legs are flexed more strongly than in the nor-
mal posture. The tarsi of the corresponding legs
are held close to each other, legs I to the front, legs
II just above the sternum, legs III just above the
epigyne and legs IV more backwards. The tarsi of
the legs are all about 0.5 cm apart from the corre-
sponding one in the web.
The female jerks fiercely every now and then
with her whole body, alternating with a slow move
from left to right, by first tightening the left legs
and then the right ones, without shifting the leg
position. Again and again she moves her abdomen
up and to the front, so that the ventral skin be-
tween the epigastric fold and the spinnerets tou-
ches a thread kept between the tips of legs III. After
that the abdomen is brought back slowly into the
horizontal position. This movement strongly re-
sembles the movement the male makes while pro-
ducing a drop of sperm. All this behaviour of the
female is continuously interrupted by spells of total
rests.
Then a clear, transparent drop appears from the
genital opening in which the white eggs are clearly
visible. She holds the drop at the top with legs II
and III. It can be seen clearly that she is pressing;
at the dorsal side of the abdomen grooves appear
in the longitudinal direction. After one minute of
pressing the drop is finished and she moves a bit
downwards. The drop appears to stick at its top to
the thread between legs III. The bottom of the
drops rest on the sternum as well as on the abdomen.
The female now begins to move her body bac-
kwards and forwards. She holds the drop with her
leg II and III, which even sink in it a little bit and
seem to stick to it. Light kneading movements are
made with legs II and III It is as if the drop is being
massaged. Thedrop becomes darker in colour. After
4 minutes and about 120 of these kneading move-
ments she stops. The drop now looks greyish and
hangs apart from the female loose on a thread,
probably the same thread that was first held by legs
III. While she holds this thread with the drop with
legs II and III, she starts to enswathe it with legs IV.
She does this in the same way as wrappinga prey. At
first she wraps the lower half of the drop. After 1
minute and 14 seconds she shifts legs II and III
upwards along the thread and starts with the upper
half. After totally 1 minute and 52 seconds she has
spun about 500 threads round the eggs and stops.
During the next 35 seconds the female bites into
the bottom of the batch, which she still holds with
legs II and III. After this she pivots round her
length axis, with the batch in her chelicerae, so that
the spot where the batch was attached to the
thread, is now below her. Then she lowers herself
along the side of the batch and with a quick stroke
of her legs she snaps the thread. Next she goes back
to a corner of the cage, her resting place, with the
egg-batch in her chelicerae (fig. 7). The drop with
the eggs still looks greyish and the eggs cannot be
distinguished any more.
From the moment of the appearance of the drop
till the withdrawal to the corner of the cage, nine
minutes have passed. Three quarters of an hour
later the drop within the strands of silk has thic-
kend a bit and the eggs are clearly visible as sepa-
rate globules. The colour of the eggs now is brow-
nish-grey, while being white when appearing in
the drop. On June 3rd, 43 days later (in cold weat-
her conditions), 63 young hatched from this egg-
batch.
Egg-batches
A. The number of batches per specimen and their
spacing in time.
Table 1 shows the dates of egg-laying by the
individual spiders, each in its own cage. All batches,
except two, were produced at night.
Data per cage.
Cage 0. - In May 1981 a male and a female were
put in this cage. During the period in which they
were together they repeatedly mated. On October
3 the male ate the female. On November 16 the
male died as well.
Cage 1. - On October 5, 1982 a female was put
in this cage. On October 26, a male was added, but
he was eaten by the female on the same day. On
April 14, 1983, again a male was brought in with
this female. Mating immediately started, after
which the male was removed. The egg-batch of
PrATEL: Pholcus phalangioides
137
Table 1. Batches per cage (= spider). Given is the date (day, month, year) and the number of eggs laid on that day
(= n).
DATE - n
30Aug81-7
24Aug83-11
[17Aug84-*]
14Aug84-17
29Aug83-30
05 Aug84-32
18Sep83-17
01Sep83-17
06Aug84-25
30Aug83-24
20Aug84-24
[11Sept84-12]
REMARKS
female eaten by male
no cocoons in 85
died in April '86
2 coc. '85 died Jul85
Female died March '85
3 coc. '85 died June '86
no cocoons in "85
died October ’85
3 cocoons in ‘85
died August '85
Also in ‘84, '85, no coc.
2 coc. '85 died Aug.'85
Number of eggs not counted.
CAGE DATE - n DATE - n
0 21May81-40 01Jul81-12
1 ie
[14May84-*]
3 ? 20Jun84-35
4 ? 17Jun82-30
21Apr83-63 30Jun83-46
[29Apr84-*] 18Jun84-48
5 n.m. n.m.
6 12Jun83-16 26Jul83-18
15May84-22 DE
7 04May83-57 26Jul83-18
16May84-37 11Jul84-26
9) 24May83-41
11 04May84-50 04Aug84-46
[date-n] = eggs not fertilized and removed by the female.
3 = Female did mate, but laid no eggs.
n.m. — Female did not mate and (thus) laid no eggs.
Female captured whenbeing adult and possibly having laid eggs before.
NB: The female in cage 7 laid eggs in 1985 on May 11, June 30 and August 17. This last batch was not fertilized.
May 14, 1984, appeared to be non-fertilized (solid,
yellowish, non globular eggs) and was removed
from the web by the female on May 19. The non-
fertilized batch of August 17, was sucked by her on
August 23, and she removed the remnants the next
day. Despite good nourishment she did not produce
eggs any more and died the 21st of April 1986.
Cage 3. - On June 18, 1984, a female was put in
this cage. This female apparently already mated, no
male was introduced.
Cage 4. - On June 8, 1984, a male and a female,
captured while they hung in the same web, were
put in this cage. On October 20, the weak male was
removed. The eggs produced on April 21, 1983,
were laid during the afternoon and the description
was given above. The eggs still fertile of August 29,
were laid 313 days after the removal of the male.
A batch laid 577 days later (on April 29, 1984)
contained infertile eggs (dark coloured) and was
removed by her on May 6. On May 22, 1984, again
a male was introduced, mating followed within a
few minutes. After recharging his palps with
sperm!), the male was removed from the cage. In
the spring of 1985 she obviously had become very
weak, while wrapping prey she inserted long bre-
aks and did not eat them any more. Part of the book
lungs appeared to have collapsed or disappeared,
while the abdomen had thickened dorsally at the
lungs level. She died on March 5, 1985.
Cage 5. - On February 8, 1984, a subadult female
was put in this cage. In February she had several
maggots for food and moulted to adulthood on
April 25. On September 4, 1984, two males were
introduced into the cage at the same time. They
mated with the female that same afternoon the one
after the other. The smaller male was frequently
chased by the larger one thus lost a leg and was
eaten by this male on October 10.
Cage 6. - On October 25, 1982, a female was put
in this cage. On June 2, 1983, a male followed and
mating immediately took place. The male died on
August 28, 1983. The egg batch of August 6, 1984,
1) A male introduced to a female, already has his palps
filled with sperm. After courtship and mating, he will
recharge his palps within a few hours.
138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
was laid 344 days after the death of the male and
contained about 40% infertile eggs. Afterwards the
female did not produce any eggs and died on Oc-
tober 4, 1985.
Cage 7. - On November 22, 1982, a female was
put in this cage. On April 8, 1983, a male was
introduced and again the mating started within a
few minutes. On May 16, 1984, she laid eggs round
12.30 hours, an event I just missed. This couple
repeatedly mated in the months of July and August.
This is the reason why the female lost her egg-
batch of August 20 on August 25, when it fell to the
bottom of the cage. This female produced three
more batches in 1985 and died early October that
year.
Cage 9. - A female was put in this cage on
October 15, 1982, followed by a male on October
26. The male was eaten by the female on Novem-
ber 2, 1982. In spite of sufficient nourishment she
laid no more eggs and died in 1985.
Cage 10. - On October 22, 1982, a subadult fe-
male was put in the cage. On May 11, 1983, this
female moulted. No male was introduced. In spite
of sufficient nourishment she laid no eggs and died
in 1985.
Cage 11. - On June 6, 1983, a female was put in
this cage. On September 2 a male was introduced
and mating started within half a minute. The eggs,
laid on September 11, 1984, appeared to be infer-
tile, despite the presence of the (weak) male; they
were removed by the female on September 13. The
male died on December 19. During 1985 the female
laid two more non-fertilized egg-batches and died
in August of that year.
If a batch with infertile eggs gets stuck in the web
somewhere, after removal by the female, she cuts
it loose again until it lies on the bottom of the cage.
Sometimes the eggs are sucked out and the remains
then removed.
As shown in table 1, the number of eggs can vary
strongly from batch to batch. The mean number of
eggs was 29.8 (n=27). The mean number of eggs
of the batches in the first column is 40.8; these are
in general the first batches of the year. In the
second column the mean is 31.0 and in the third
one, the last batches of the year, the mean is 19.9.
Some of the batches are not used (non-fertilized,
difficult to place) for calculating these averages.
The infertile eggs in batches otherwise normal
have been counted.
B. Influence of food on egg-production.
Table 2 shows the time interval between a meal
of protein rich prey, such as queens of Lasius niger
Table 2. The laying of eggs after consumption of a Lasius
niger queen.
Date of cons. Cage Date of egg Days
L. niger number production later
19 Aug83 1 24Aug83 5
19Aug83 4 29Aug83 10
20Aug83 6 01Sept83 12
19Aug83 2 30Aug83 11
29Sept83 all none —
30Jul84 1 17Aug84 18
30Jul84 3 14Aug84 15
30Jul84 4 05 Aug84 6
30Jul84 6 06Aug84 7
30Jul84 TE 20Aug84 21
30Jul84 11 04Aug84 5
* The female in cage 7 still had an egg-batch when con-
suming a L. niger.
about to fly out, and the production of eggs: an
average of 10 days (the female from cage 7 not
counted, see table 2). If also other prey animals
(smaller flies, midges, Tipulids) are taken into ac-
count the mean number of days between consump-
tion and egg-production was 12.6. In February all
spiders were fed with one or more maggots or
freshly emerged flies.
The observations described above can be sum-
marized as follows.
In my house Pholcus phalangioides produces
eggs from the end of April to the end of September
and almost always during the night. Only in the
case a female is well-fed and mates for the first
time, eggs are laid soon after the mating. If a female
is less well-fed the eggs are laid only when enough
food becomes available.
If a first mating takes place towards the end of
the year, the eggs are laid in the following spring.
The female of P. phalangioides appears to be able
to store sperm so that fertilization can still take
place 344 days after mating. The amount of sperm
received during copulation is sufficient for three
egg-batches. After a few days batches only contai-
ning infertile eggs are remove from the web by the
female.
Food consumption appears to stimulate the
production of eggs. Females are able to lay nine
egg-batches during the three years they are adult.
The mean number of eggs per batch is 29.8. The
number of eggs in the first batch of the year usually
is larger than in the second batch and lowest in the
third batch of the year. Females that did not mate
are absolutely unable to produce any eggs.
PLATEL:
Pholcus phalangioides
139
Table 3. Duration of the development in days of the eggs and the second stage (prenymphs).
A B C D E
4 3 30Jun83 16 5
7 2 26Jul83 18 6
6 D 26Jul83 18 6
3 2 14Aug84 19 10
6 5 06Aug84 20 7
il 2 OáAug8á 20 8
4 7 OSAug84 21 7
7 6 20Aug84 23 12
6 1 12Jun83 23 5
4 6 18Jun84 24 10
7 5 11Jul84 24 7
3 1 20Jun84 24 9
A B C D E
1 l 24Aug83 26 8
4 4 29Aug83 28 8
9 l 24May83 28 4
7 3 30Aug83 30 8
6 3 01Sep83 il 9
7 4 16May84 36 8
6 4 15May84 36 7
7 1 04May83 37 8
1 l 04May84 39 8
5 1 18Sep84 41 14
4 2 21Apr83 43 7
A = Cage number; B = egg-batch number of the female concerned; C = date of egg production; D = duration of the
development of the eggs in days; E = duration of the development of the prenymphs, in days, till the second moult.
Duration of development of the eggs
All egg-batches studied were laid in 1983 and
1984 by eight females (table 3). The time needed
for development in days, from the moment of
laying until the hatching of the young, can vary
considerably (table 3).
Figure 1 shows the duration of the development
of 23 egg-batches put against the mean outdoor
temperature during that period. It shows to be
strongly temperature-dependent. The mean dura-
tion of the development is 27.2 days (S.D. 8.0) with
a mean outdoor temperature over this period of
ISO RENS D N27):
Female behaviour while carrying eggs
It was observed regularly that an egg batch car-
rying female attached it to the web in order to be
able to eat, mate or to clean herself, but only a few
days after laying the eggs. The female therefore
attaches a thread to the web and brings it, with the
aid of both legs III, slowly to the batch, the tarsi
touching each other. She then takes the batch in her
legs I and Ill and attaches a thread to it by touching
the batch a few times with her spinnerets at the
point where it is kept by legs III. Then she looses
the batch, and by now it is suspended from the web.
A prey can be wrapped by the female while she
carries the batch in her chelicerae, but more often
she attaches the batch to the web first. If a female
carries an egg-batch a male has to exert himself
more than usually to obtain a mating, which in
most cases meets with success. When a male tries
to insert his palps while the female still holds the
eggs in her chelicerae, he does not succeed. When
after elaborate courtship by the male, the female is
willing to mate (the epigyneal region has swollen),
she first attaches the batch to the web, mates and
then returns to her eggs immediately afterwards.
She snaps the supporting lines with her legs Ill and
takes them back into her chelicerae again, someti-
mes after first having cleaned herself.
When disturbed the female always immediately
returns to her eggs and takes them, in case of a
serious disturbance, back in her chelicerae, or resu-
mes after a short while, in case of a less serious
interference, eating prey, cleaning, but never ma-
ting.
4 6 8 10 12 14
days
Fig. 1. Duration of the development of eggs in relation
to the outdoor temperature. | = indicates the average;
__ = linear regression-line: y = —0.327x + 24351; r =
0977 p <<0:001;n'=723.
140 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
egg tooth
Figs. 2—5. Pholcus phalangioides. Fig. 2. Young in ruptured chorion. (Drawn from a slide.) Fig. 3. Young stuck in
the remainder of the prelarval integument. Hairs no longer covered by the prelarval integument stand erect, others,
still covered shine through the skin and lie flat. Fig. 4. Part of a leg (femur-tibia joint) still within the prelarval
integument. (Drawn from a slide.) Fig. 5. Tarsus of a leg still within the prelarval integument. The tarsal claws can
be seen within it. (Drawn from a slide.)
Dispersal of the young
Some 10 to 15 days before hatching a breach in
the chorion is visible (fig. 8), leading from the
frontal side, just below the clypeus of the young
spider, backwards to the side of the abdomen which
is folded forwards against the sternum (fig. 2). The
eyes which are already discernible, and part of the
carapace and the palps as well as the coxae of the
legs show through the crack (fig. 9).
About this time the female starts to loosen the
batch a bit, which is necessary because there is an
increase in the diameter of the batch (approxima-
tely 0.5 mm, as was later measured with other
eggs). She does this by biting through the threads
round the eggs. She attaches the batch to the web
as previously described, holds it with legs II and III
with which she turns the batch in order to reach
other threads. She then picks up the threads very
carefully with the cheliceral fangs without touching
the eggs and bites them through or pulls them free.
Afterwards she takes the batch in her chelicerae
again.
This behaviour was observed 13 days before hat-
ching at the earliest. If she has once started this
behaviour, she shows it regularly and with incre-
asing frequency and duration at each bout, as the
time of hatching approaches. Towards the time the
young hatch, one can perceive a regular indenting
of their carapace, just behind the eyes. After a while
the spiderling starts stretching its legs and con-
PLATEL:
15 20 25 30 35 40 45
days
Fig. 6. Number of days between hatching and the second
moult (= the first moult outside the chorion), in relation
to the outdoor temperature. | = gives the average; __ =
linear regression-line; y = —1.055x + 24.334 r = —0.876
: p << 0.001; n = 22.
tinues to do so until it has freed itself and has
reached the outside of the batch. Soon all the
spiderlings are on the outside.
Often the female still holds the batch in her
chelicerae and after a while she then has a cluster
of spiderlings hanging on her jaws (fig. 10). It also
occurs that the female attaches the batch to the web
when the young are about to emerge or just after
the emergence of the first young. Shortly after
hatching, till some hours hereafter, the young
crawl along the webstrands and disperse just above
the remnants of the batch, within a circle with a
diameter of about 5 cm, depending on the number
of spiderlings. When some spiderlings have remai-
ned in the batch, the female is seen to brush thé
batch, carefully but quickly, with legs II and III and
sometimes also legs IV (fig. 12). She also goes on
biting the threads, besides carefully evading the
bodies of the young spiders. She may be engaged
with this behaviour up to three days, with intervals.
The young that are still caught in the strands as
well as the undeveloped eggs are sucked out, so that
only a skin remains, after which she removes the
remnants from the web.
As an experiment some batches were filched
from the female, at an early stage and not without
some effort. They were suspended on a tiny needle
Pholcus phalangioides 141
in a jar. The eggs developed quite normally, with
the exception of a few that were damaged. During
hatching it was obvious that those young had great
difficulties to free themselves from the spinnings
(fig. 11). Even a single thread could give insur-
mountable troubles. In general the young which
were more on the outside succeeded most of the
time in freeing themselves after some hours of
hard labour. As to the young more to the centre of
the batch this effort took much longer. Some of
those young did not manage to free themselves
even after three to four days of continuous struggle.
These young died. Spiderlings near the outside, but
caught in the threads with one or more legs, can
sometimes free themselves during the first moult
outside the chorion. When this does not meet with
success they die after all.
Summarizing we may state that 10 to 16 days
before hatching the chorions break. From 13 days
onwards before hatching the female loosens the
batch more and more by cutting the threads around
the eggs. The young emerge quickly, whereby the
stragglers are being helped by the mother: she
brushes the batch with her legs and cuts threads.
The batch is then still held with the chelicerae or
attached to the web. Young that stuck are sucked.
In an experimental situation, with no female pre-
sent, the young clearly have more difficulties free-
ing themselves.
Development of young
By closer investigation part of the young proves
to have died in the remainder of the batch while
still sitting partly in the chorion. It reveals that the
part that still sticks, the legs, is enveloped by a very
thin skin. The hairs on the free part of the leg stand
erect, while the hairs on the part still covered lie
completely flat (fig. 3). The skin that envelopes the
legs, contains neither hairs nor tarsal claws and is
not segmented either. Through this very transpa-
rent skin the characteristics of the next stage can
be seen, i.e. the new skin still wrinkled, with hairs
and tarsal claws (figs. 4 and 5). Hence it appears
that the young that comes free nog only frees itself
from the chorion but also from the first prelarval
integument. Some of the young that died, stuck in
this moult (fig. 3). The prelarval integument is so
thin and transparent, that nothing recognizable
can be found after hatching.
The most remarkable characteristics of the
emerging spiderlings are given below.
Mobility: The spiders move little, disperse inside
a small region in the web around the mother and
move away when touched.
142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
10
Figs. 7—11. Pholcus phalangioides. Fig. 7. Female with eggs. Fig. 8. Female with loosened batch of eggs, a few days
before the dispersal of the young. Fig. 9. Prelarvae after the rupture of the chorion; in the two prelarvae on the left
the blackish egg-tooth on the palp is visible. Fig. 10. Female with freshly hatched spiderlings. Fig. 11. A spiderling
struggling to get free.
Cuticle: Not pigmented and with fewer hairs but less than with the nymph. One trichobothrium
than with the nymph. on each metatarsus. The foot claws are toothed.
Legs: all joints are present. Hairs are present, Chelicerae: The fang is present and looks nor-
PLATEL:
Pholcus phalangioides
143
Fig. 12. Female of Pholcus phlangioides assisting the spiderlings to free themselves from the webstrands.
mal, but the poison-duct is not clearly visible (a
poison-duct is not necessary; they still live on their
yolk reserves).
Spinnerets: Spinnerets and silk glands are pre-
sent. The young are able to spin threads.
Eyes: The eyes look normal, but are surrounded
by a brown pigmented area and not by a black area
as in older spiders. The eyes are not elevated and
lie flat on the carapace.
I have never seen prenymphs spin threads them-
selves, if they had webstrands of their mother at
their disposal (normal situation). I only observed
the latter with spiderlings that hatched from eggs
suspended from a needle. One day after emergence
the spiderlings are able to spin threads themselves.
These lines are double and thin and are sometimes
drawn out of the spinnerets with the aid of the
tarsal claws of legs IV. They are able to attach these
lines efficiently, and the fast movements of the
spinnerets are then clearly visible. When seriously
disturbed they partly drop on a thread spun by
themselves. Later they climb back along the same
thread. The prenymphs in the mother’s web do not
yet show the characteristic rapid whirling move-
ments of this family. They pull themselves up a bit
between their legs at most, while the mother in-
deed vehemently whirls. In the first nymphal stage
the young do sometimes show this behaviour.
After a period of a few days the young sit very
still. When they place the tarsi of the legs close to
or against each other the second moult begins.
The time that passes between hatching and this
second moult is, as with the development of the
eggs, strongly temperature-dependent (fig. 6). On
the average moulting starts 7.9 days after hatching
(S.D.=2.3; n=22; table 3), with a mean outdoor
temperature during this period of 16.0 °C
(SD 27)
A period of 15 days was noted with young that
developed in a batch taken away from the female
(November 8, 1983).
DISCUSSION
Egg laying, a case study
Egg laying with Pholcids has been described ear-
lier by Pokrowsky (1899) for Pholcus opilionoides
and by Maughan (1978) for Ph. muralicola. Pok-
rowsky possibly had Ph. phalangioides in mind. In
the past century these two names were use as syn-
onyms (Simon, 1866). Van Hasselt (1870) also
mentioned Ph. opilionoides while in view of the
size of his specimens he clearly had Ph. phalan-
gioides in mind. Nowadays both names are used for
different species.
144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
In a few lines Maughan (1978) describes egg
laying with Ph. mwralicola, a species resembling
opilionoides and phalangioides but being a little bit
smaller. Both Pokrowsky and Maughan describe
the female’s posture before laying eggs as a resting
posture. They do not mention the tarsi placed close
to each other which is not the case in the resting
posture. Observations in the literature on egg
laying with Ph. phalangioides are possible lacking
because most of the eggs are not laid in the daytime
and the proper laying is only of short duration.
Montgomery (1903) “…watched pregnant females
for long intervals in the hope of seeing it...”, but
without success. With Ph. muralicola eggs were
extruded by pulsating movements of the abdomen
(Maughan, 1978). Pokrowsky (1899) describes the
female making strange movements with the abdo-
men extruding the eggs with the aid of abdominal
muscles. These movements may correspond with
the female’s movements observed by me. The ac-
tion of the abdominal muscles can be seen: the
grooves in the longitudinal direction which arise
when squeezing. Pholcids do not possess dorsoven-
tral muscles (Kaston, 1981) so that these grooves
must be caused by the longitudinal muscles.
The colour of the eggs when laid is in accordance
with the observations of Pokrowsky and Maughan.
With Ph. Muralicola eggs became brown or black
as development proceeded (Maughan), while Po-
krowsky only describes that the eggs become dry
and lose their lustre after a few minutes. Maughan
(1978) writes nothing about the presence of a fluid
during laying, while Pokrowsky (1899) only men-
tions that they are “made wet by a fluid”. However,
I clearly observed a drop in which the eggs appe-
ared. The presence of such a viscid drop was des-
cribed with more species. In those cases it was
observed that the viscosity of the fluid is such that
the egg mass largely retains its globular shape. This
drop also contains sperms (Gertsch, 1979) or pos-
sibly sperms (Montgomery, 1903). The viscosity of
the drop with Ph. phalangioides is so strong that
this, without being wrapped in silk, keeps hanging
from a thread all by itself, so that the spider can
start throwing silk around it. Assuming that the
drop contained sperms, it might be possible that
the function of the 4 minutes kneading movements
by the female is to produce a better contact of the
sperms with the eggs. However, it is also possible
that the female checks the viscosity of the drop and
waits until this is strong enough by drying up. In
doing so the female can leave the drop hanging on
a line without the danger of its flowing out and
next she can start wrapping it. The former assump-
tion does not exclude the other. The kneading
movements were not observed by Pokrowsky and
Maughan. Mention is made of the help of legs
while laying (legs III, according to Maughan; with
one of her legs, according to Pokrowsky). Po-
krowsky already saw some threads around the eggs,
before the female started enswathing them. In
whatever way this is possible is not clear to me.
Pokrowsky’s observation about the female enswa-
thing the eggs with her front pair of legs must be
wrong.
The snapping of the thread on which the egg-
batch hangs with the aid of the legs was observed
more frequently by me with females that tempora-
rily hung the batch in the web and took it back in
the chelicerae after eating, mating or cleaning.
Montgomery (1903) also described this behaviour.
It is clear by now that Pholcus enswathes the eggs
with silk. Earlier it was believed that the eggs were
merely glued together (Bonizzi, 1869; Becker,
1892), but in what way can the female carry the
eggs without damaging them? However, in those
days there were authors who did mention a thin
silk covering around the eggs (among others: van
Hasselt, 1870). Later it was confirmed by more
authors (Montgomery, 1903; Bonnet, 1930b; Chry-
santhus, 1950; Wiehle, 1953; Bristowe, 1941, 1958;
Gertsch, 1979; Kaston, 1981). Carrying the egg-
batch in the chelicerae, probably to protect it, is a
thing that is mostly mentioned with the descrip-
tions of this species. The silk threads around the
eggs are very thin and probably this is the reason
why they were not observed by some authors. Yet
the number calculated of about 500 lines around the
eggs is more than expected.
Egg-batches
The dates for the first batch given in literature
are not widely spread. Chrysanthus (1950) saw the
first egg-batch on April 26. Montgomery (1903)
and Bristowe indicated the month of May for this.
The last batches were seen in August (Bristowe,
1958). Montgomery (1903) even saw a female lay
eggs in November. However, Schaefer (1976) ob-
served Ph. phalangioides laying eggs the whole
year through, with a maximum in May. The popu-
lation investigated by him lived in cellars, where
the temperature, in winter, too, did not fall below
20 °C. Schaefer calculated a zero point of develop-
ment with about 14 °C and also mentioned that
females are able either to mate or lay eggs and
young are not able to hatch at temperatures below
10 °C. The lower temperatures in winter causes
the periodicity found in this and other researches.
Bonnet (1930b) in this aspect also mentions; “that
spiders do not moult at temperatures lower than
PLATEL:
13 °C, one can assume that this temperature at
least is necessary, because tissues have to be formed
which also applies to the development of the young
in the eggs”.
Bonnet (1930b) and Schaefer (1976) mention
that a condition for the female to lay eggs is that
she must have mated, which also appears from this
research. Mating does not necessarily have taken
place just before laying. Bristowe (1958) already
stated that a female is able to produce eggs in May
or June, after having mated in the previous sum-
mer. Schaefer (1976) mentions a period of 17 to 36
weeks. Now it was even observed that fertile eggs
can be laid after a period of 11 months after mating.
Some of the eggs in this batch were infertile, pro-
bably because of a shortage of (vital) semen. Eggs
are laid under good conditions indeed, rather soon
after the first mating. This was also observed by
Bonnet (1930b) and Chrysanthus (1950). In that
case the period between mating and laying is
roughly in accordance with the two weeks Seitz
(cited in Foelix, 1979) mentions and which are
necessary for the increase of the volume of the eggs
in the second stage, which starts after mating. An
increase in diameter of the female’s abdomen was
not noticed by me and was not measured either.
This again is in accordance with the observations
of Gerhardt (1921). Montgomery (1903) and Ka-
ston (1981) wrote that females did not produce
more than three cocoons. He probably means: not
more than three cocoons per year. It appears that
females are able to produce at least six batches with
a maximum of nine.
Egg-batches falling on the bottom of the cage
only contain infertile eggs in all cases and are re-
moved by the female herself, sometimes after they
were first sucked by the female. Van Hasselt (1870)
saw up to two times the complete disappearance of
a cocoon and then supposed them to be sucked by
the female. Montgomery (1903) noticed that from
the 12 cocoons observed by him 4 had dropped and
did not hatch. He writes that these cocoons drop-
ped by accident when hung in the web. Of the 32
batches observed by me, only one has dropped by
accident, apart from the 4 clearly nonfertilized
ones, which were removed by the female herself.
This was the last egg-batch - in this research - of
the female in cage 7. She had to hang the batch in
the web many times because of the many matings.
On the fifth day after laying she fished up a remain-
der of an earlier prey in her chelicerae instead of
the batch which lay on the bottom. An egg-batch
does nog simply fall out of the chelicerae. If one
wants to take away the batch from a female, a
relatively great force has to be used. It takes a lot
Pholcus phalangioides 145
of trouble and it will not go without destroying a
large number of eggs. Bonizzi (1869) and Montgo-
mery (1903) already stated that the female will not
quit the egg-batch even if treated roughly. I assume
that the 4 cocoons “lost” by Montgomery's spiders,
contained infertile eggs and were removed by the
female herself, because none of them produced any
young. The egg-batch lost by the female in cage 7,
which could be saved, did produce young. Yet a
dozen eggs did not survive the fall and treatment
afterwards.
A connection between the moment of food con-
sumption and the time of egg production is not
mentioned in the literature.
The number of eggs for Ph. phalangioides given
in the literature lies between 13 and 60 (Bonnet,
1930b; Chrysanthus, 1950; Wiehle, 1953; Kaston,
1981). But these are incidental observations. Only
Schaefer (1976) gives a mean number of 33.2 eggs
per batch (n = 16, 28-46 eggs).
The diameter of the batches was not measured
by me. Locket & Millidge (1951) give about 5 mm
for this, and Wiehle (1953) 4 mm. Kaston (1981)
even gives the diameter in relation to the number
of eggs: a batch of 4.7 mm contained 29 eggs; an
other of 3.6 mm had 25 eggs. The size of the eggs
was also given by him: 1.2 to 1.3 mm long and 0.94
to 1.2 mm thick. On other cocoons of
Ph. phalangioides I ascertained later the following
diameters and egg numbers: diameter 3.6 mm — 36
eggs; 4.5 mm - 42 eggs; 5.4 mm - 60 eggs. The
greater part of the time the first egg-batches are
bigger than the next, as with many more spider
species that lay eggs more than once (Chrysanthus,
1950; Gertsch, 1979; Foelix, 1979). In addition
Gertsch mentions that some of the later eggs may
be infertile, owing to the exhaustion of the semen
supply stored in the receptacles, and perhaps also
to its gradual loss of viability (see also Montgo-
mery, 1908). With Ph. phalangioides the latter
only seldom happens because the females nearly
always are susceptible to the males, this in contrast
with the observations of Montgomery (1903) and
Gerhardt (1921, on Ph. opilionoides). If with
Ph. phalangioides the male stays away after a co-
pulation, the phenomenon of infertile eggs does
occur, but only if afterwards several batches are
produced. In general the number of eggs in spring
is again larger than the number in the autumn of
the previous year, but smaller than in the first
batch laid by the female. But as is already menti-
oned the number of eggs also depends on the sup-
ply of food so that differences can also arise because
of this.
146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Duration of development of the eggs
It is only Schaefer (1976) that gives the duration
of the development of the eggs in relation to the
temperature: 18.1 days with 23 °C (n= 8) and 54.7
days with 16 °C (n= 6). This is in accordance with
the data of this research. Other authors only men-
tion the duration of the development observed by
them, which most of the time lies between two and
three weeks (Bonizzi, 1869; Montgomery, 1903;
Bristowe, 1958). Chrysanthus (1950) mentions a
very long duration of 52 days for eggs laid in spring.
Female behaviour while carrying eggs
According to most authors the females of
Ph. phalangioides carry the eggs in their chelicerae
from laying up to, or up to and including, the
emergence of the young (van Hasselt, 1870; Bec-
ker, 1892; Wiehle, 1953; Vachon, 1965; van Ka-
twijk, 1976; Gertsch, 1979; Kaston, 1981; Nen-
twig, 1985). It was also observed that the female
hung the batch in the web, in order to eat, clean
herself (Bonizzi, 1869; Montgomery, 1903; Chry-
santhus, 1950), or to copulate with an insistent
male (Bristowe, 1958). My observations about the
way in which the female hangs the eggs in the web
and looses them again, is in accordance with Mont-
gomery’s observations (1903).
Dispersal of the young
Bonizzi (1869) described the female trying to break
the egg shells with her mandibles one day before
hatching. He probably observed the breaking of the
threads around the eggs. Bonizzi did not observe
these threads. For the rest Bonizzi is the only aut-
hor mentioning this kind of behaviour before the
emerging of the young. However, Ph. phalangioi-
des starts biting these threads already sooner, pro-
bably as soon as the chorions rupture by an increase
of volume of the embryo (Bonnet, 1930a; Gertsch,
1979; Foelix, 1979) and with the aid of the sharp
egg- teeth at the basis of the palps (Vachon, 1965;
Gertsch, 1979). This biting of the threads also
occurs with other spiders, particularly wolf-spiders
(among others: Montgomery, 1903; Bonnet,
1930a; Foelix, 1979). The fact that females still bite
threads after the emergence of the first young, was
also observed by Montgomery (1903). This aid
appeared to be indispensable for many young.
There are clearly individual differences, for some of
the females have repeatedly worse results than
others.
The time during which the young, after hat-
ching, stay together in a cluster before dispersing
in the web to some extent, can vary from some
hours (Bonizzi, 1869; Montgomery, 1903) up to some
days (Becker, 1892); Chrysanthus, 1950), as also
became clear from this research.
Development of the young
The development and morphology of the young
of Ph. phalangioides after the rupture of the cho-
rion, the presence of a prelarva and a prenymph,
are excellently described by Vachon (1965). Emerit
(1984 and in a pers. comm., 1985) also emphasizes
the presence of a prelarva. The characteristics sum-
med up by me clearly show that the spiderling
described is a prenymph (Vachon, 1957, 1965;
Emerit, 1984). Canard (1984) uses a different ter-
minology in which these differences with Vachon
in relation to Ph. phalangioides are given in table
4 (drawn up by Canard, 1985 in a pers. comm.). His
terminology shows the different stages in develop-
ment better, but it is difficult to see when the first
period (pullus) ends (with the rupture of the first
integument). Accurate observation of the young
still partially present in the chorion is necessary.
The duration between the rupture of the chorion
and the emergence of the prenymph amounts from
5 tot 10 days according to Vachon (1965), but in
this research from 10 to 15 days. Other authors do
not mentioned this period, because in general it
was assumed that the young did only moult for the
first time a few days after hatching.
In the literature the duration of this prenymphal
stage is given as 6 to 7 days (Bonnet, 1930b); 10
days (Vachon, 1965) or 7 to 14 days (Bristowe,
1958). Schaefer (1976) again gives the duration in
relation to the temperature: 17.3 days with a con-
stant temperature of 16 °C (n = 21) and 7.9 days
with a constant temperature of 23 °C (n= 26). In
this period the young live on their yolk reserves
and are not able to take food themselves (Vachon,
1965), so that the assertion of Bonizzi (1869), that
a female spun a fly for them (“...when the more
robust of the young animals ran to suck the insects
thus prepared for them...) must be questioned.
The possibility exists that these “more robust of
the young animals” were prenymphs already moul-
ted. A first nymph sometimes carefully tries to suck
a prey spun by the female together with her. Ho-
wever, the young was repeatedly spun by the fe-
male with the prey. A young that was noticed when
sharing in the profit, was simply swept away with
a leg each time.
ACKNOWLEDGEMENTS
I would like to thank Dr. Peter van Helsdingen
for his helpful advice and critical reading of the
manuscript. I also like to thank Mr. Van Gils and
Katy van Helsdingen for the translation.
PLATEL:
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Katwijk, W. van, 1976. Spinnen van Nederland. - Rotter-
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Montgomery, T. H., 1903. Studies on the habits of spid-
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Montgomery, T. H., 1908. The sex ratio and the cocoon-
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Schaefer, M., 1976. Experimentelle Untersuchungen zum
Jahreszyklus und zur Überwinterung von Spinnen
(Araneida). - Zoologische Jahrbücher, Systematik
103: 127-289. |
Simon, E., 1866. Monographie des espèces européennes
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ment postembryonnaire des araignées. Première note.
Généralités et nomenclature des stades. - Bulletin de
la Société Zoologique de France 82: 337-354.
Vachon, M., 1965. Contribution à l'étude du developpe-
ment postembryonnaire des araignées. Troisième
note. Pholcus phalangioides (Fussl.) (Pholcidae). —
Bulletin de la Société Zoologique de France 90 (5/6):
607-620.
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Received: 24 August 1988
Accepted: 30 December 1988
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THE TAXONOMIC POSITION OF THE PACHYNEURINI
(CHALCIDOIDEA, PTEROMALIDAE) AS JUDGED BY
CHARACTERISTICS OF COURTSHIP BEHAVIOUR
by
J. VAN DEN ASSEM & M. J. GIJSWIJT
ABSTRACT
Assem, J. van den & M. J. Gijswijt, 1989. The taxonomic position of the Pachyneurini
(Chalcidoidea, Pteromalidae) as judged by characteristics of courtship behaviour. — Tijdschrift
voor Entomologie 132: 149-154, tabs 1-2. [ISSN 0040-7496]. Published 3 July 1989.
Characteristics of courtship behaviour seem to be of considerable interest from a comparative
point of view. A syndrome of behavioural features was used for separating a group of pteromalids
(viz. the Pachyneurini) from the Pteromalinae, a subfamily to which they were formerly assigned.
Key words. - Pteromalidae; Pachyneurini; taxonomy; behaviour.
Dr. J. van den Assem, Zoological Laboratory, Division of Ethology, University of Leiden, P.O.
Box 9516, 2300 RA Leiden, The Netherlands
INTRODUCTION
Current classification of the Chalcidoidea is still
far from what must be the ultimate aim: an arran-
gement of taxa reflecting the group’s phylogeny.
Comparative studies of chalcidoid mating behav-
iour can make a contribution to this end (Gauld
1986) because they provide, in principle, independ-
ent tests of classifications that were based on mor-
phological criteria only (the usual practice). Fea-
tures of behaviour may serve as characters on
different levels. Such features are often species-
characteristic; sibling species in particular may be
identified more easily on characteristics of mating
behaviour than of morphology (e.g. Van den As-
sem & Povel 1973). Just as well, behaviour ele-
ments can serve to separate genera or families (e.g.
Van den Assem et al. 1982b). Actually, the use of
behavioural features as criteria of classification is of
a long standing (e.g. the classification of Anatidae
— ducks and relatives -: Heinroth 1910, Lorenz
1941). In the entomological practice severe limita-
tions abound since living specimens are not usually
the kind of material at the disposal of taxonomists
and, moreover, far from all species will “behave”
under laboratory conditions. However, this is no
excuse; phylogenetic classifications require sup-
portive evidence from various biological sources.
Those components of courtship repertoires
which function as intra-specific signals qualify as
149
useful characters in the first place. They are sup-
posed to have acquired their more or less fixed
quality during evolution in a process called ritual-
isation. Fixation means minimal variation in per-
formance — a necessary quality of unambiguous
signals and, likewise, of reliable diagnostic charac-
ters. Courtship signals produced by chalcidoid
males are, at least partly, of a chemical and/or of
an acoustic nature, and cannot be recorded or an-
alysed without sophisticated equipment (Van den
Assem 1986). On the other hand, signal emission
usually coincides with overt movements of one or
more limbs, and these movements can be described
and quantified in a simple way (in terms of number
of occurrences or of durations; e.g. Barrass 1960;
Martin & Bateson 1986).
Previous studies of chalcidoid mating behaviour
have corroborated current classifications (e.g. Ko-
gan & Legner 1970, Van den Assem & Povel 1973),
or suggested changes (e.g. Van den Assem et al.
1982a, Dahms 1984, In den Bosch & Van den As-
sem 1986). In the present paper we want to point
out an apparent discrepancy, and suggest a change.
MATERIAL AND METHOD
Parasitized hosts were collected in the field and
individual parasitoids were isolated prior to emer-
gence, as far as possible, to obtain virgin speci-
mens. (Females of several species are known to
150 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
mate only once in a lifetime; in some species they
appear no longer attractive to males following ma-
ting. Obviously, virgin specimens are necessary for
studying courtship and mating behaviour in such
cases). A list of the material that was used for the
observations mentioned in this paper is given in
the appendix.
All observations were made with a low-power
binocular microscope. Prior to the observations, a
male and a female specimen were introduced into
a small, perspex cell that was closed with a glass
cover slip. The cell’s diameter (ca 2.5 cm) corre-
sponded to the visual field of the microscope. Mag-
nifications of 10 or 15 X proved to be sufficient.
Many sequences of courtship and mating behaviour
were recorded on video tape.
COURTSHIP BEHAVIOUR OF PTEROMALINAE
Among taxonomists there is consensus that
several large families of Chalcidoidea (e.g. Encyr-
tidae, Fulophidae, Furytomidae) represent monop-
hyletic entities while others do not (e.g. Pteroma-
lidae, Torymidae). Many lower ranking taxa appear
to be non-holophyletic assemblages: the Pte-
romalinae, a sub-family of Pteromalidae, may serve
as an example. Clearly, the Pteromalinae are a
‘rest’ group. Positive morphological criteria for
separating Pteromalinae from alternative groups
do not exist. Unifying behavioural characteristics
do not exist either. Yet, within this group, several
clusters of distinctly related genera can be defined,
some of which were already recognized by
Thomson (1878). Similar clusters are apparent in
the checklist of Chalcidoidea by Boutek & Graham
(1978): genera are arranged in a non-random or-
der, suggestive of varying degrees of relatedness
(never explicitly stated by the authors). Examples
are: Psilonotus-Anogmus-Mesopolobus; Nasonia-
Trichomalopsis (= Eupteromalus); Caenasis-Cect-
dostiba-Hobbya-Ablaxia-Aggelma. Details of
courtship and mating procedures give support to
these assumed relationships (unpubl. data).
For the Pteromalinae as a whole, unifying featu-
res of courtship and mating behaviour cannot be
defined. Yet, for what seems to be a large majority,
an assemblage that we refer to as the Pteromalinae
s.s., such features do exist. Earlier (Van den Assem
1974), this group has been characterized as “those
pteromalids in which the male takes up a frontal
position on the female for courtship (by placing his
front feet on her head) and produces a sequence of
motor coordinations in which the antennae are
moved synchronously as each other's mirror image,
the general direction of these movements being up
and down; the female indicates her readiness to
copulate by a special antennal signal (she draws her
flagellae tightly to her head capsule) and maintains
a frozen posture for some time”. This definition —
which was based on observations of 14 species —
may still serve, although the importance of mouth-
part extrusions (see below) as taxonomic criteria
was not appreciated at the time.
The elements of courtship and mating which
characterize the Pteromalinae s.s. are mentioned
below. Actually, it is the syndrome of elements that
is characteristic; taken separately, alle features can
be found in many other groups as well. Our obser-
vations now refer to 37 species (mentioned in the
appendix) belonging to the 20 genera enumerated
in table 1. We assume that many more can be added
to the Pteromalinae s.s. once courtship and mating
procedures are known. Our prediction is that the
rules which refer to general characteristics of mo-
vements, temporal structure, and frontal position
of the male (see below) will apply to all near-
relatives of the genera mentioned in table 1.
Characteristic behaviour patterns
Courtship position. - The male courts from
a frontal position on top of the female, with its fo-
re tarsi placed on the female’s head. The precise
placement differs between species (e.g. laterally
on the female’s eyes, on its nape, near the base
of its scapes, etc.). By necessity, a male has to back
up from its courtship position before it can copu-
late.
Antennal movements. — The male’s antennae,
whatever the precise motor coordinations, move as
each other’s mirror image. The details of these
movements, and the corresponding temporal pat-
terns, are usually characteristic for a species or for
a group of species. Several kinds of movements are
involved; one of them is most conspicuous: the
antennal sweep, a swift up-and-down large ampli-
tude movement, and performed just before or syn-
chronously with a mouthpart extrusion (below).
Extrusion of the mouthparts. - The male extru-
des (lowers) its mouthparts periodically in a con-
spicuous way (as if ‘vomitting ). Extrusions may
be combined with head-nodding movements. If so,
an extrusion coincides with the upstroke of a nod.
Female receptivity signal. - At the onset of se-
xual receptivity (i.e. coinciding with the exposure
of the genital orifice) the female lowers its anten-
nae (previously held horizontally). It draws the
flagellae tightly in to the front of its head (flagellae
pointing downwards).
VAN DEN ASSEM & GIJSWIJT:
Table 1. Genera of Pteromalinae s.s.; courtship and ma-
ting procedures were observed in one or more species
Anisopteromalus Ruschka, 1912
Anogmus Förster, 1856
Caenasis Förster, 1856
Cecidostiba Thomson, 1878
Dibrachys Förster, 1856
Dinarmus Thomson, 1878
Hobbya Delucchi, 1957
Hypopteromalus Ashmead, 1900
Lariophagus Crawford, 1909
Meraporus Walker, 1834
Mesopolobus Westwood, 1833
Muscidifurax Girault & Sanders, 1910
Nasonia Ashmead, 1903
Peridesmia Förster, 1856
Psilonotus Walker, 1834
Pteromalus Swederus, 1795 (including the subgenus
Habrocytus Thomson, 1878)
Stenomalina Ghesquière, 1946
Trichomalopsis Crawford, 1913
Trichomalus Thomson, 1878
Trychnosoma Graham, 1957
Duration of genital contact. - In Pteromalinae
s.s. the duration of genital contact (at 20 °C) is
relatively long, ranging between 10 secs (in e.g.
Nasonia) and ca 1 minute (in e.g. Lariophagus).
An experimental analysis of the displays of a few
species of Pteromalinae s.s. has revealed (Van den
Assem 1986) that mouthpart extrusions are asso-
ciated with the release of chemical stimuli (pher-
omones). Head-noddings probably enhance the ef-
fects of pheromone release. There are (slight)
differences in the precise motor coordinations be-
tween clusters of related genera (e.g. the nodding
movements of Anogmus - Mesopolobus species
differ from those of Nasonia vitripennis and rela-
tives). Several clusters include genera with nodding
and non-nodding species. Also, within certain ge-
nera (e.g. Habrocytus) species may differ in the
extent of nodding. Apparently, nodding has been
“invented” a number of times, which means that
not all nods are truly homologous.
COURTSHIP BEHAVIOUR OF PACHYNEURINI
Courtship displays in one section of Pteromali-
nae (i.e. the Pachyneurini sensu Ashmead, 1904)
do not correspond to those of the Pteromalinae s.s.
We believe that the differences are of a fundamen-
tal kind (see Discussion), and we suggest to sepa-
rate the Pachyneurini from the Pteromalinae and
preferably raise it to sub-family rank, the Pachy-
neurinae. Ours is not the only argument: Copland
& King (1972) found consistent differences in the
Pachyneurini 151
structure of the female reproductive system of
Pachyneuron versus other species of Pteromalinae.
Moreover, Boutek and Graham (1978) placed the
Pachyneurini at the end of the Pteromalinae in
their checklist, not because they believed it to be
the most apomorphic section, but because it was
unclear where to place it at all (Boutek, pers.
comm.). Earlier, Boutek (1961) has suggested that
the (then newly described) genus Vrestovia was
close to Synedrus Graham, 1956, Ablaxia Delucchi,
1957 or Caenasis Förster, 1856, but this suggestion
was dropped later. Features of courtship would not
support such a relationship either: Caenasis males
court in the way of Pteromalinae s.s.; males of the
two Vrestovia species we have seen do not.
Judged by general characteristics of their mating
behaviour, the Pachyneurini represent a natural
unit. The diagnostic syndrome comprises the fol-
lowing features:
Characteristic behaviour patterns
Courtship position. - The male’s fore tarsi are
placed at the frontal edge of the female’s pronotum
(on its shoulders”) or on top of its thorax. In
Pachycrepoideus vindemmiae the placement is va-
riable: sometimes on the female’s thorax, someti-
mes on its head (but never stereotypical on the
head).
Phase shifts. - There are periodic shifts in the
orientation of the courting male with respect to the
female below. Foreward-backward movements
with the entire body occur which coincide with
bending-stretching movements of the fore legs
while the fore tarsi remain in the same place
throughout. (These postural changes are the equi-
valents of the periodic low phase-high phase shifts
described by Van den Assem et al. 1982b.)
Antennal movements. - All displays include mo-
vements with the antennae; however, not in all
phases are the antennae moved as each others
mirror image. There is no antennal sweep of the
kind seen in Pteromalinae s.s., but there is a more
distinct forward-backward component in the an-
tennal motions.
Mouthpart movements by the male. - Forward-
backward movements are always present, but of an
inconspicuous kind which is easily overlooked.
There are no real extrusions which suggest “lic-
king” actions, as in the Pteromalinae s.s., and the
motor patterns involved differ greatly. (In the
Pteromalinae s.s. mouthpart extrusion is a prom-
inent element of a male’s display.)
152 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Position of the female antennae. — During
courtship, females point their antennae more or
less vertically upward, forming an acute figure V.
This posture never occurs in female Pteromalinae
S.S.
Mouthparts female. — Typically, females hold
their mandibles agape as soon as they are mounted.
Female antennal signal and duration of genital
contact. — Antennal movements made by the fe-
male at the onset of receptivity occur in some
species, but judged from direct observations it
seems doubtful that they serve a signal function as
found in Pteromalinae s.s. Backing up by courting
males may involve considerable delays. Antennal
movements by the female were absent in Vrestovia
and Pachyneuron species (antennae remain in the
upright position throughout) and of a variable
quality in Pachycrepoideus, from upright to hori-
zontal or below. In Toxeumorpha the antennae go
into a low position. But even when low, the ensuing
posture is less “complete” than in the Pteromali-
nae s.s. (the flagellae are not tightly drawn in to the
front of the head) (table 2).
The duration of genital contact varies between
species. In most it is very short (a matter of a few
seconds only); in Toxexmorpha contacts may last
for over a minute.
DISCUSSION
The courtship procedures of species of Pteroma-
linae s.s. have many features in common. Species
of Pachyneurini were found to be different in this
respect, not just in a matter of degree but in a
fundamental way because they lack a number of
what we consider to be synapomorphic features.
By examining the display behaviour of many
species, it is possible to uncover phylogenetic
trends (i.e. successively more advanced combina-
tions of behavioural traits). Similar trends can be
traced in several large families of Chalcidoidea,
suggestive of parallel developments. Two of these
are relevant in the context of this paper: the switch
of the male’s courtship position from the rear to
the front, in connection with the development of
a (secondary) receptivity signal, and the omission
of periodic shifts in the position of the courting
male relative to the female (Van den Assem 1986).
In this respect the Pteromalinae s.s. qualify as
synapomorphic: the position of the courting male
has moved to the extreme front, the male’s head is
just above the female’s antennae and the male
releases a necessary, receptivity inducing stimulus
periodically — probably a product of its mandibular
glands — by means of extruding its mouthparts.
Head-noddings enhance the effectiveness of stimu-
Table 2. Genera and species (in alphabetical order)
which were observed to deviate from the courtship and
mating procedures of the Pteromalinae s.s.
Pachycrepoideus Ashmead, 1904; Pachyneuron Walker,
1833; Toxeumorpha Girault, 1915; Vrestovia Boutek,
1961.
Pachycrepoideus vindemmiae (Rondani): Curepe, Trini-
dad, from pupae of an antomid fly September 1971;
Hanoi, Vietnam, from pupae of Drosophila spec. Lab-
oratory culture on D. melanogaster.
Pachyneuron muscarum (L.): Vleuten, The Netherlands,
parasite of Leptomastix dactylopu Girault, on Plano-
coccus citri in a glasshouse; May 1988.
Pachyneuron planiscuta Thomson: Leiden, The Nether-
lands, emerged from reed stalks, host unknown,
March 1974.
Toxeumorpha nigricola (Ferriere): Natal, Rep S Africa,
from pupae of Zaprionius spec. in tomato fields, 1980.
Laboratory stock on D. melanogaster.
Vrestovia fidenas (Walker). Zuidwijk, Wassenaar The
Netherlands, on pupae of Drosophila spec; April
1987.
Vrestovia spec: Amherst. Mass USA, on pupae of Dros-
ophila spec.; October 1979. (Bou£ek in litt).
lus release. Females have developed a secondary
receptivity signal which is a sufficient stimulus for
the male to stop courting and back up (Van den
Assem & Jachmann 1982). Phase-shifts in the
male’s position are absent, males court in a low-
phase position throughout. Pachyneurini give evi-
dence of a less advanced condition. The position of
the courting male is less extreme; males change
their posture periodically during a display (without
shifting the position of the forefeet); there is no
evidence for pheromonal stimulation by way of
specialized movements with the mouthparts. A
secondary antennal signal is absent, or, where an-
tennal movements occur, evidence that males take
them as a cue for switching to copulatory behaviour
is missing. Actually, we often observed a delay
between the moment of overt receptivety (the
moment the female exposes its genital orifice) and
the male backing up, and it seems to be longer, on
average, than in the Pteromalinae s.s., although in
this group as well males are variable in this respect.
To delay backing up to copulate may have serious
consequences for a courting male because sneaking
competitors may take precedence and mate first
(which always means: inseminate more success-
fully). In this respect the procedures of Pachyneuron
muscarum are of interest. Males produce a single
courtship cycle, back up immediately and attempt to
copulate. If the female is not yet receptive then the
frontal courtship position is taken up again, a new
cycle follows, etc. A similar organization of the
courtship display has been observed in Pteromalus
VAN DEN ASSEM & GIJSWIJT:
puparum, it was hypothesized that it might be
understood as an adaptation to a high level of
competition (Van den Assem 1974). Pteromalus is
a parasitoid of the chrysalids of butterflies from
which hundreds of wasps may emerge more or less
synchronously. The same explanation may hold for
the Pachyneuron muscarum display. Moreover,
males hold their wings low while backing up, in a
cape-like fashion (as do some tetrastichids in an
attempt to ward off competitors) thus providing an
additional argument. This rapid-pendulum type of
temporal organization will prolong the time requi-
red to induce receptivity in a female but this loss
will probably be more than evened out by losing a
copulation to a competitor less often.
For Pteromalus, it was argued that the pendulum
procedures were probably not an original feature
(Pteromalus females have a fully developed anten-
nal signal), but for Pachyneuron it may be diffe-
rent. Courting males provide conspecific females
with stimuli that may induce sexual receptivity. In
probably genuinely primitive groups (such as Ce-
rocephalinae) males stay in a caudal position
throughout, and a successful courter can thus per-
ceive the onset of receptivity immediately (it recei-
ves a tactile stimulus, the effect of the female’s
abdomen-raising). Pachyneurini males are small
enough, and the position on the female is enough
to the front to make a direct perception of recep-
tivity unlikely. The most effective strategy would
then be to make an inspection on the spot repe-
atedly, which is what Pachyneuron muscarum ma-
les do. However, males of other species, which
likewise lack a secondary signal, do produce a se-
quence of cycles before backing up. We have no idea
on what cues they might act.
We have argued that the Pachyneurini should be
separated from the Pteromalinae and be made a
sub-family in its own right. Judged by characteri-
stics of display behaviour, the Pteromalidae com-
prise an array of subfamilies which combine origi-
nal character states (e.g. Cerocephalinae,
Spalanginae), highly derived character states
(Pteromalinae s.s.), or intermediate conditions
(e.g. Miscogastrinae, Asaphinae). The Pachyneu-
rini belong to the latter category but they do not fit
into one of the existing subfamilies. The conspicu-
ous position of the females’ antennae offers no cues
(similar positions are observed in many groups
throughout the Chalcidoidae). The movements of
the males’ mouthparts do not provide cues either,
nor do characteristics of the respective repertoires.
For the time being the Pachyneurini should be kept
apart from other groups, awaiting a more profound
analysis of mutual relationships.
Pachyneurini 153
ACKNOWLEDGEMENTS
Many colleagues sent us specimens for observa-
tions. Their help is greatly appreciated. We are
indebted to Drs. Z. Boutek (London) and M. W. R.
de V. Graham (Oxford) for identifications and dis-
cussions.
REFERENCES
Assem, J. van den, 1974. Male courtship patterns and
female receptivity signal of Pteromalidae, with a con-
sideration of some evolutionary trends and a comment
on the taxonomic position of Pachycrepoideus vinde-
miae. - Netherlands Journal of Zoology 24: 253-278.
Assem, J. van den, 1986. Mating behaviour in parasitic
wasps. — In: Insect Parasitoids (J. K. Waage & D.
Greathead eds). 13th Symposium Royal Entomologi-
cal Society London: 137-167.
Assem, J. van den, H. A.J. in den Bosch & E. Prooy, 1982a.
Melittobia courtship behaviour: a comparative study
of the evolution of a display. - Netherlands Journal of
Zoology 32: 427-471.
Assem, J. van den, M. J. Gijswijt & B. K. Nübel, 1982b.
Characteristics of courtship and mating behaviour
used as classificatory criteria in Eulophidae — Tetra-
stichinae, with special reference to the genus
Tetrastichus. — Tijdschrift voor Entomologie 125:
205-220.
Assem, J. van den & F Jachmann, 1982. The coevolution
of receptivity signalling and body-size dimorphism in
the Chalcidoidea. - Behaviour 80: 96-105.
Assem, J. van den & G. D. E. Povel, 1973. Courtship
behaviour of some Muscidifurax species (Hym. Pter-
omalidae): a possible example of a recently evolved
ethological isolating mechanism. - Netherlands Jour-
nal of Zoology 23: 465-487.
Barrass, R., 1960. The courtship behaviour of
Mormoniella vitripennis. - Behaviour 15: 185-209.
Barrass, R. 1976. Courtship behaviour in Meraporus gra-
minicola and other Pteromalinae. - Physiological Ent-
omology 1: 77-81.
Bosch, H. A. J. in den & J. van den Assem, 1986. The
taxonomic position of Aceratoneuromyia granularis
(Hym. Eulophidae), as judged by characteristics of its
courtship behaviour. - Systematic Entomology 11: 19-
237
Bouéek, Z., 1961. Beiträge zur Kenntnis der Pteromali-
den-Fauna von Mitteleuropa, mit Beschreibungen
neuer Arten und Gattungen (Hym). - Acta Entomo-
logica Musei Nationalis Pragae 34: 55-95.
Boutek, Z. & M. W. R. de V. Graham, 1978. Hymenoptera.
— Checklist of British insects (G. S. Kloet & W. D.
Hincks eds) 4: 67-110.
Copland, M. J. W., & P. E. King, 1972. The structure of
the female reproductive system in the Pteromalidae.
- Entomologist 105: 7-96.
Dahms, E. C., 1984. Revision of the genus Melzttobia
(Chalcidoidea Eulophidae) with the description of se-
ven new species. - Memoirs of the Queensland Mu-
seum 21: 271-336.
Gauld, I. D., 1986. Taxonomy, its limitations and its role
154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
in understanding parasitoid biology. — In: Insect Parasi-
toids (J. K. Waage & D. Greathead, eds). 13th Sympo-
sium Royal Entomologial Society London: 137-167.
Heinroth, O., 1910. Beiträge zur Biologie, namentlich
Ethologie und Psychologie der Anatiden. - Internati-
onal Congress of Ornithology 5: 589-702.
Kogan, M. & E. F Legner, 1970. A biosystematic revision
of the genus Muscidifurax (Hym. Pteromalidae) with
descriptions of four new species. — Canadian Entomo-
logist 102: 1268-1290.
Lorenz, K., 1941. Vergleichende Bewegungsstudien an
Anatiden. — Journal fiir Ornithologie 89: 194-294.
Martin, P. & P. Bateson, 1986. Measuring behaviour. An
introductory guide: 1-200. — Cambridge University
Press.
Miller, M. C., R. White & C. Smith, 1973. Courtship
behaviour of Peridesmia discus. - Entomophaga 17:
20-25.
Thomson, C. G., 1878. Hymenoptera Scandinaviae V. —
Lund.
APPENDIX
Anisopteromalus calandrae (Howard): Savannah Geor-
gia USA, on weevils in wheat stored in warehouse;
January 1971. Laboratory culture on Sitophilus.
Anogmus hohenheimensis (Ratzeburg): Oberstdorf
BRD, on cecidomyids in cones of Picea abies; January
1979.
Anogmus piceae (Ruschka): Kiental Switzerland, on
idem; August 1978.
Anogmus strobilorum Thomson: Oberstdorf BRD, on
idem; February 1977.
Anogmus vala (Walker): Oberstdorf BRD, on idem; Fe-
bruary 1977.
Caenasis lauta (Walker): Leiden The Netherlands, from
galls of Cyntps divisa; February 1986.
Cecidostiba semifascia (Walker): Mt Ventoux France,
from galls on Quercus spec.; August 1978.
Dibrachys boarmiae (Walker): Meyendel Wassenaar The
Netherlands, parasitic on a dipterous parasite of an
earwig inside stems of Asparagus spec.; April 1978;
laboratory culture on Calliphora.
Dibrachys cavus (Walker): Wageningen The Nether-
lands, from Apantheles pupae on Pieris spec.; March
1972.
Dinarmus basalis (Rondani): Slough England, from a
laboratory culture on Callosobruchus chinensis in be-
ans; May 1981.
Habrocytus bedeguaris Thomson: Bentheim BRD, from
galls on Rosa spec.; April 1974.
Habrocytus elevatus (Walker): Oberstdorf BRD, on try-
petids in flowerheads of Centaurea spec.; February
1978; Oegstgeest The Netherlands, on idem in flower-
heads of Arctia spec.; June 1978.
Habrocytus sequester (Walker): Wijster The Nether-
lands, from seedpods of Linaria vulgaris; September
1977.
Hobbya stenonota (Ratzeburg): Mt Ventoux France,
from galls on Quercus spec.; August 1978.
Hypopteromalus tabacum Ashmead: McNeil in litt 1974.
Lariophagus distinguendus (Forster): Leiden The Net-
herlands, obtained as an infection of our laboratory
culture of Sitophilus granarius on wheat.
Meraporus graminicola Walker: quoted by Barrass
(1976).
Mesopolobus rhabdophagae (Graham): Wassenaar The
Netherlands, from galls on Salix repens; July 1983.
Mesopolobus dubius (Walker): Antequera Spain, from
galls of Plagiotrochus on Quercus spec.; March 1978.
Mesopolobus mediterraneus (Mayr): Antequera Spain,
idem.
Mesopolobus fasciiventris Westwood: Bunde The Net-
herlands, from galls of Pediaspis aceris on Acer; June
1974.
Muscidifurax raptor Girault & Sanders Riverside Califor-
nia USA, obtained from laboratory cultures on Musca
domestica, details quoted in Kogan & Legner (1970).
Muscidifurax raptorellus Kogan & Legner: idem.
Muscidifurax uniraptor Kogan & Legner: idem; Septem-
ber 1975.
Muscidifurax zaraptor Kogan & Legner: idem, as M.
raptor.
Nasonta vitripennis (Walker): Leiden The Netherlands,
on pupae of Calliphora spec; August 1971.
Nasonta spec. nov.: Rochester NY USA, from bird nests
in nestboxes; August 1987 (this species has long-
winged males).
Peridesmia discus (Walker): in Miller, White & Smith
(1973), and Miller in litt, 1974.
Psilonotus achaeus Walker: Meyendel Wassenaar The
Netherlands, from Semudobia galls on Betula spec.;
October 1976; Oberstdorf BRD, idem; March 1978;
's Graveland The Netherlands, idem; April 1985.
Psilonotus adamas Walker: Meyendel Wassenaar The
Netherlands, idem; August 1976.
Pteromalus puparum (L): Ales France, from chrysalids of
Pieris spec.; August 1971.
Pteromalus venustus Walker: Lethbridge Alberta Canada,
from pupae of Megachile rotundata; November 1974.
Stenomalina liparae (Walker): Leiden The Netherlands,
from galls of Lipara lucens in Phragmites spec. Janu-
ary 1971.
Trichomalopsis (Eupteromalus) micropterus (Linde-
mann): Oegstgeest The Netherlands, on drosophilids
in apple orchard, August 1986.
Trichomalopsis (Eupteromalus) spec. Riverside Califor-
nia USA, from a laboratory culture on Drosophila
spec. from locally collected material.
Trichomalopsis (Eupteromalus) tigasis (Walker): De
Lemmer The Netherlands, collected as adults on a
window; October 1973.
Trychnosoma punctipleura (Thomson): Oberstdorf
BRD, from cecidomyids in cones of Picea abies; Febru-
ary 1977.
Received: September 1988
Accepted: 9 January 1989
PHYLLOGOMPHOIDES INDICATRIX, A NEW DRAGONFLY FROM
MEXICO (ODONATA: GOMPHIDAE)
by
JEAN BELLE
ABSTRACT
Belle, J., 1989. Phyllogomphoides indicatrix, a new dragonfly from Mexico (Odonata: Gom-
phidae). — Tijdschrift voor Entomologie 132: 155—157, figs 1-7. [ISSN 0040-7496]. Published
3 July 1989.
Phyllogomphoides indicatrix spec. nov. from Mexico (state of Chiapas, Ixhuatan) is described
and illustrated after a unique male. Corrections on some earlier papers are provided as well.
Key words. — Gomphidae; Phyllogomphoides new species; Mexico.
J. Belle, Onder de Beumkes 35, 6883 HC Velp, The Netherlands.
INTRODUCTION
In the State of Chiapas, Mexico, Dr. Rosser W.
Garrison collected a male of Phyllogomphoides
which he recognized as belonging to a new species.
He kindly offered me to describe it and for this
privilege I herewith wish to thank him very much.
In this paper I present a description and illus-
trations of this dragonfly under the name
Phyllogomphoides indicatrix, the specific name be-
ing suggested by Dr. Garrison himself.
The new taxon belongs to the semicircularis
species group (cf. Belle 1984), having the superior
anal appendages semicircular-forcipate. However,
it is very distinct from all other members of the
group in the form of the anterior genital hamule
which in complexity is nearest to that of Phyllo-
gomphoides pacificus (Selys). The thoracic colour
pattern resembles that of Phyllogomphoides sua-
sillus Donnelly by the well-developed (first and
second) pale antehumeral stripes and the (three)
pale lateral stripes.
Phyllogomphoides indicatrix spec. nov.
(figs. 1-7)
Holotype @: Mexico, State of Chiapas, Ixhuatan
(Teapa-Chiapa de Corzo Rd, 400 m), 26 September 1983,
R. W. Garrison leg. (National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C.).
Description of male. — Measurements: total
length 60 mm; abdomen 45 mm (incl. appendages
155
3.1 mm); hind wing 36 mm; costal edge of ptero-
stigma in fore wing 4.5 mm.
Head: face brown with conspicuous pale (=
bright yellow) markings as follows: labrum with
two large pale lateral spots, base of mandibles and
genae pale, anteclypeus pale, and postclypeus with
two large pale lateral spots. Upper part of frons
basally dark brown, anteriorly with a broad bluish
grey band which is narrowed in middle. Vertex
dark brown, the depressed central area bright yel-
low. Occipital plate bright yellow with dark brown
borders.
Prothorax: dark brown but middle lobe with
a yellow spot on either lateral side and a yellow
middorsal twin-spot.
Pterothorax: dark brown with bright yellow an-
tehumeral stripes and greenish yellow lateral
stripes; its colour pattern shaped as shown in fig.
1
SS
Legs: femora dark brown but inner sides of first
pair of femora greenish yellow. Tibiae, tarsi and
claws black.
Wings: slightly brown tinged, its venation black
but frontal margin of costae with a fine yellow
line. Pterostigma reddish dark brown. Basal sub-
costal cross-vein present. Discoidal triangles, sub-
triangles and supratriangles three-celled but sub-
triangle and supratriangle in left fore wing four-
celled. Nodal index 14:22—22:14/17:16—15:15.
Second primary antenodal cross-vein the seventh.
Intermedian cross-veins 12—10/7—8. Hind wings
156 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
width four-celled anal triangle, a three-celled anal
loop, six (left) and five (right) paranal cells, a
single row of five cells in second anal interspace but
in left hind wing with an extra small paranal cell,
and area posterior to Cu2 four (proximal) to five
(distal) cells wide.
Abdomen: dark colours on basal segments dark
brown, on other segments black. Pale colours on
basal segments greenish yellow, on other segments
yellow. Segment 1 pale on middorsum and lateral
sides. Segment 2 with pale auricles, a pale postero-
lateral spot, pale ventral tergal margins, and a pale
middorsal stripe which is rather broad and parallel-
side on the basal two-thirds and very narrow on the
apical third. Segments 3 and 4 with an interrupted
pale middorsal line and pale basal side spots. Seg-
ments 5 and 6 with pale basal spots on middorsum
and on sides. Segment 7 pale for its basal third and
along ventral tergal margins. Segment 8 pale on
proximal half of lateral dilatations; the greatest
width of these foliations about 0.3 mm. Segments
9 and 10 without pale markings. Superior appen-
dages brown-yellow but black on the basal fourth
and at tips of projections. Inferior appendage
brown. Accessory genitalia and anal appendages
shaped as shown in figs. 2—7.
ERRATA
I take the opportunity of publishing four correc-
tions of errors, noticed in some of my previous
papers.
1964. A new dragon fly of the genus Rhodopygia.
— Studies on the fauna of Suriname and
other Guyanas 7, p. 51. The legend of fig. 23
seminal vesicle should be “glans of penis”.
Figs. 1—7. Phyllogomphoides indicatrix spec. nov., holotype &: 1, thoracic colour pattern; 2, tenth abdominal segment
and appendages, dorsal view; 3, accessory genitalia, ventral view; 4, the same, right profile view; 5, anterior genital
hamule, right profile view; 6, the same, left profile view; 7, apical segments of abdomen and anal appendages, left
profile view.
BELLE: Phyllogomphoides 157
1972. Further studies on South American Gom-
phidae (Odonata). — Tijdschrift voor Ent-
omologie 115, p. 223. The legend of fig. 15
dentata should read “edentata”
1973. A revision of the New World genus
Progomphus Selys, 1854 (Anisoptera, Gom-
phidae). — Odonatologica 2, table 1 (p.
198). The type location of anomalus spec.
nov., MNHW should be “MZM”.
1988. A synopsis of the species Phyllocycla Cal-
vert, with descriptions of four new taxa and
a key to the genera of Neotropical Gomphi-
dae (Odonata, Gomphidae). — Tijdschrift
voor Entomologie 131, p. 96. The numbers
of the figures 74 and 75 should be interchan-
ged.
REFERENCE
Belle, J., 1984. A synopsis of the South American species
of Phyllogomphoides, with a key and descriptions of
three new taxa (Odonata, Gomphidae). — Tijdschrift
voor Entomologie 127: 79— 100, figs. 1—40.
Received: 11 October 1988
Accepted: 20 January 1989
EPIGOMPHUS CORNICULATUS, A NEW DRAGONFLY FROM
COSTA RICA (ODONATA: GOMPHIDAE)
by
JEAN BELLE
ABSTRACT
Belle, J., 1989. Epigomphus corniculatus, a new dragonfly from Costa Rica (Odonata: Gom-
phidae). — Tijdschrift voor Entomologie 132: 158-160, figs. 1-6. [ISSN 0040-7496]. Published
3 July 1989.
Epigomphus corniculatus is described from Costa Rica. The nearest relative of this species
is E. armatus Ris.
Key words. — Epigomphus; taxonomy; Costa Rica.
Dr. J. Belle, Onder de Beumkes 35, 6883 HC Velp, The Netherlands.
INTRODUCTION
During his field work in Costa Rica, Mr Carlos
Esquivel of the Universitdad Nacional at Heredia
(Costa Rica) collected a male and two females of
an undescribed species of Epigomphus. He kindly
placed this material at my disposal, for description
and reporting, and for this privilege I herewith
wish to thank him very much.
The nearest relative is Epigomphus armatus Ris,
1918. Ris’ figure 93 is somewhat misleading. It
shows the male inferior anal appendage in an ob-
lique ventral view from the rear. In a strictly ven-
tral (or dorsal) view the posterior margin of the
inferior appendage is widely V-shaped with stout
branches (fig. 1). The male of the new taxon is
readily distinguished by the very widely U-shaped
posterior margin of the inferior appendage (fig.
2) while the corresponding female differs in hav-
ing the postocellar tubercles produced backward
to a pair of well-developed “horns” (fig. 4).
All figures have been drawn with the camera
lucida by the author. The details were added by
freehand.
Epigomphus corniculatus spec. nov.
(figs. 2-6)
Material. — Prov. Limón: Suretka, 23 April 1988, 16 1
2 (in cop.; holotype and allotype, respectively), 1 9
(somewhat teneral; paratype). All three specimens pre-
served dry in clear plastic envelopes. The holotype and
allotype are deposited in the Florida State Collection of
Arthropods (FSCA), Gainesville. The paratype is in the
Museo de Insectos, Universidad de Costa Rica, San José.
158
Description of the male holotype (abdomen
broken between segments 2-4).
Measurements. Total length 54 mm; abdomen
(incl. app.) 41.5 mm; hind wing 34 mm; costal edge
of pterostigma of fore wing 3.3 mm.
Head. Dark brown with pale markings as fol-
lows. Genae green; labrum with a symmetric pair
of large, round, grey-green lateral spots; postcly-
peus with grey-green facial lobes; superior surface
of frons with an anterior grey band that is narrowly
interrupted in middle; vertex yellowish brown be-
tween each lateral ocellus and compound eye. Dor-
sal surface of occiput with two shallow transversely
elongated concavities occupying each of its lateral
two-fifths, and there is a development of an occip-
ital ridge which, however, is not fringed with hairs.
Prothorax. Dark brown, the middle lobe green-
yellow on sides and on middorsum.
Pterothorax. Dark brown with pale (= greyish
green) markings. Pale mesothoracic “half collar”
interrupted in middle and not connected with first
pale antehumeral stripe. Second pale antehumeral
stripe narrow but complete. Pale mesepimeral and
metepisternal stripes well-developed. Posterior to
the dark metapleural stripe the metepimeron is
pale except for an obscure brown area on center
line. Pectus pale.
Legs. Femora brown, the third pair blackish at
knees, the first and second pair blackish on outer
sides. Tibiae, tarsi and claws blackish brown. An-
tero-inferior row of spines on third tibiae consist-
ing of 10-11 modified spines, that of first and se-
BELLE:
Epigomphus corniculatus 159
Fig. 1. Epigomphus armatus Ris, 1918. Tenth abdominal segment and anal appendages of male, ventral view. — Figs
2-6. Epigomphus corniculatus spec. nov.: 2, tenth abdominal segment and anal appendages of male holotype, ventral
view; 3, vulvar lamina of female allotype, ventral view; 4, head of female allotype, dorsal view; 5, apical abdominal
segments and anal appendages of male holotype, dorsal view; 6, the same, left profile view.
cond joints of third tarsi also with modified spines.
Abdomen. Dark brown with pale (= green-yel-
low) markings on segments 1 to 8 as follows. Seg-
ments | and 2 pale on lateral sides and with a pale
middorsal stripe. Segments 3 to 6 with a pale basal
spot on each lateral side. Segments 3 and 4 with a
pale basal middorsal stripe. Segments 5 and 6 with
a small pale basal middorsal spot, the one on seg-
ment 6 much smaller than that on segment 5.
Segment 7 pale on basal two-thirds portion. Seg-
ment 8 with a small pale basal spot on each lateral
side. Abdomen narrow on segments 3 to 6, beco-
ming wider successively on apex of segment 7 and
on segments 8 to 10, being on segment 10 five
times as wide as on base of segment 7 (fig. 5).
Superior appendages somewhat longer than infe-
rior appendage, widely distended but not enough to
give a dorsal view of the inferior appendage. Apex
of each superior appendage rounded and crenula-
ted, the outer margin angled. Inferior appendage
with two widely separated slender branches, each
branch with a bifid apex, the mesal tooth small (fig.
2). Dorsal surface of inferior appendage with a
posterior pair of submedian humps and more ba-
sally with a pair of stout acute teeth.
Description of the female allotype (left hind
wing broken off, left anal appendage broken away).
Measurements. Total length 57 mm; abdomen
(incl. app.) 43 mm; hind wing 39 mm; costal edge
of pterostigma of fore wing 4.0 mm.
Coloration resembling that of male holotype but
pale colour on superior surface of frons consisting
of two widely separated anterior spots. Dark colour
of vertex and top of head reddish brown. Dark
colour of prothorax brown. Spines on outer row of
third femora widely spaced and in length about half
the diameter of femur. Abdominal segment 8 wit-
hout pale basal side spots. Abdominal segment 7
pale for slightly more than its basal half. Abdomen
160 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
becoming successively narrower on apical seg-
ments. Lamina supra-analis slightly shorter than
anal appendages, the latters about three-quarters
the length of segment 10. Vulvar lamina subtrian-
gular, reaching to a point about halfway the ninth
sternum, medially cleft for its apical three-fifths
(fig. 3).
Except for the pair of long post-ocellar “horns”
the most striking mating adaptations resemble
those of Epigomphus armatus as described by Cal-
vert, 1920 (page 342). There is a very deep pit on
the rear of the head behind each compound eye and
the top of the head (occiput) has a symmetric pair
of transversely elongated, submedian, dorsal pits.
The posterior margin of the occiput has a small but
distinct median excision (fig. 4) contrary to
Epigomphus armatus which has the rear margin
medially slightly concave. Each compound eye has
a striking large superior ocular scar near the angle
of the mesal margin of the eyes caused by the tips
of the branches of the male inferior appendage
during the copulation (the female paratype has not
such a pair of ocular scars apparently because, seen
the teneral condition of the specimen, it has not yet
paired).
REFERENCES
Calvert, P. P., 1920. The Costa Rican species of Epigom-
phus and their mutual mating adaptations (Odonata).
— Transactions of the American Entomological So-
ciety 46: 323-354, pls 13-15.
Ris, F, 1918. Libellen (Odonata) aus der Region der
amerikanischen Kordilleren von Costarica bis Cata-
marca. — Archiv für Naturgeschichte 82 (9): 1-197,
tabs 1, 2.
Received: 6 April 1989
Accepted: 12 April 1989
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G. van Rossem, The genus Cryptus Fabricius, 1804 of the Canary Islands (Hymenoptera,
Ichinéumontdae) stee sn un ee ART CT LPS SR en
J. Belle, A revision of the New World genus Newraeschna Hagen, 1867 (Odonata: Aesh-
TGA EG) Mr ee Ain a ae eye ONE or MG Gb oc ooccoocvo
P. Kanaar, A new Saprinus from Irian Jaya (Coleoptera: Histeridae).................
Rectification [to Platel, The egg laying and larval development of Pholcus phalangoides]
Published 1 December 1989
ISSN 0040-7496
ALEXEY DIAKONOFF, 1907-1989
The editors of the Tijdschrift voor Entomologie
recently received the sad news that our former
editor, Dr A. N. Diakonoff, passed away at the age
of 82 on September the 20th, 1989. We share our
feelings of sadness with the Dutch Entomological
Society, of which he has been an active member for
very many years, and with his family, that so much
took part in his life as an entomologist.
Alexey Diakonoff was born in Saint Petersburg
(now Leningrad) in 1907 and experienced all the
miseries of a country going through a revolution.
In 1923, by a circuitous way, he was able to join his
parents who, according to plan, had reached the
Netherlands East Indies. There he finished his
elementary education and left for Amsterdam for
a university study in biology. As a trained entomol-
ogist, his thesis on Indo-Malayan Tortricidae com-
pleted, he returned to Java in 1939 to become an
entomologist at a research station of the sugar
plantations and industries. Just when he had ob-
tained, in 1941, a position of entomologist at the
Zoological Museum at Bogor, Java, fate, in the
shape of World War II, prisoner of war camps and
deportation, prevented him from starting a scien-
tific museum career. In 1945 he returned to The
Netherlands for recuperation, most of the time
staying at the Leiden Museum and working in the
Lepidoptera collection. In 1947 he returned to Bo-
gor, but not for long. The political situation
changed very rapidly and the future looked uncer-
tain. In 1951 he returned to The Netherlands to
become curator of Lepidoptera at the Rijksmuseum
van Natuurlijke Historie in Leiden.
Back in The Netherlands he at once became an
active member of the Dutch Entomological Society.
Already at the summer meeting in June 1953 it was
reported that Diakonoff had joined the editorial
board of the Society publications. In fact the last
issue of 1952 of the “Tijdschrift” and the wrapper
of the completed volume carried his name. And
what is more, the journal changed in several as-
pects, in external appearance as well as in the
internal lay-out. First of all the “Tijdschrift” was
freed of the reports of the society s meetings and
the Annual Report, which from then on would be
published in the "Entomologische Berichten”. The
“Tijdschrift voor Entomologie” thus became a
scientific journal in its purest form. At the same
time the lay-out was modified. Distinct page head-
ings with the journal’s name on one page and a
running title of the article on the opposite page
161
Dr. Alexey Diakonoff in his working room, 1982.
Photo: E. L. M. van Esch.
were introduced. The size of the pages became
larger, a new letter-type (Times Roman) was se-
lected, indeed the whole format of the journal was
modernized. If one opens the 1953 volume one
recognizes the familiar make-up of recent decades,
which indeed stood its ground up till now. Each
page is proof of the strong preference Diakonoff
had for symmetry in the text. The only change
made was the recent shift to a two-column distri-
bution of the text, and that only for economical
reasons and not on aesthetic grounds, and after
Diakonoff had left the editorial board.
In 1953 there also appears an emblem on the
wrapper and title-page: Lycaena dispar batavus
Oberthiir, the Dutch subspecies of the Large
Copper, on top of a strongly enlarged egg of the
same species, surrounded by the name of the So-
ciety. The emblem was modified after the vignette
used at the occasion of the 9th International Con-
gress of Entomology at Amsterdam (1951). Al-
162 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
though we are not absolutely certain, we have a
strong feeling that Diakonoff had a firm hand in all
these developments, could convince the board with
his aesthetic arguments and obtained a mandate to
reshape the “Tijdschrift”.
In the early sixties Diakonoff plead for the crea-
tion of another series of publications, the "Mono-
grafieën van de Nederlandse Entomologische
Vereniging”. They were intended to serve as a
medium for longer articles, which otherwise would
surpass the size of a yearly volume of the Tijd-
schrift. In 1964 the first volume was published,
while in 1968 a monograph on the Dutch Tortri-
cidae, written by him in cooperation with Count
Bentinck, appeared as volume 3 of this series.
Many other papers by his hand appeared in our
journal, amidst a total of about 250 scientific pap-
ers, in which he - amongst others — described
hundreds of species and genera of Microlepidop-
tera, in particular from the Indonesian archipelago.
In 1970, at the occasion of the 125th anniversary
of the society, Diakonoff was granted the honorary
membership of the society because of his long and
highly appreciated efforts to improve upon the
international standing of the society s publications.
In 1974 he left editorship after more than twenty
years of service, because he wanted to devote all his
time to scientific work. His successors on the ed-
itorial board call themselves happy to have been
trained by him, to have received his lessons in the
printing trade, to have benefited from his long
experience. Modesty kept him from ever criticizing
our work on the later volumes, but we know he
followed all developments. He couldn't set aside
his interest in “his” Tijdschrift after so many years
of involvement, or could he?
Only two weeks before his death, Alexey Diako-
noff sent us a book review for publication in this
journal. Below we publish this review, apparently
one of Diakonoff's last manuscripts, as a homage
to this great predecessor of us.
P. J van Helsdingen (former editor)
E. J. van Nieukerken (editor)
BOOK REVIEW
Medvedev, G. S. (ed. ), 1987 [1988]. Keys to the Insects
of the European part of the U. S. S. R., vol. iv, Lepi-
doptera, part 1. - Published for the United States
Dept. of Agriculture and the National Sciences Foun-
dation, Washington, by Amerind Publ. Co., New
Delhi, xxvi + 991 pp, 578 figs. — Translated from
Russian by D. R. Sharma. - [Distributed by E. J. Brill,
Leiden, Netherlands, price US $ 97.50]
This book covers the most archaic groups of
Lepidoptera: the suborders Micro- and Macrojuga-
tae and several minor families of the suborder
Frenatae, including the economically so important
Tortricidae, 17 families in all, the other 16 being:
Micropterygidae, Eriocraniidae, Hepialidae, Nep-
ticulidae, Opostegidae, Tischeriidae, Heliozelidae,
Incurvariidae, Adelidae, Psychidae, Heterogynidae,
Limacodidae, Zygaenidae, Sesiidae, Cossidae and
Atychiidae. The species composition of the Euro-
pean part of the USSR is almost completely re-
viewed (about 1200 species). Identification keys are
(mostly) at the species level. Illustrations of the
genitalia of both sexes are given for all species,
with notes on morphology, biology and distribu-
tion of families and genera. Phenology and food
plants of most species are discussed. Finally bibli-
ography and indices of insect and hostplant names
conclude this book.
This important translation represents a part of
the Russian "Opredelitel” (=Keys) series, written
by leading specialists of the insect-fauna of the
European part of the USSR. The original series is
very popular among international Lepidopterists,
in spite of the barrier of the language. I even know
some colleagues, who taught themselves Russian,
in order to be able to read the text. Others acquired
the Russian edition, only to study the illustrations
of wing neuration and genitalia, and often also of
the wing pattern. Now that the language problem
is overcome, the translated edition will fill a most
important gap in the western bibliography on Le-
pidoptera.
[A. Diakonoff]
NEW DRAGONFLIES (ODONATA) FROM COSTA RICA
by
STEPHEN J. BROOKS
ABSTRACT
Brooks, S. J., 1989. New dragonflies (Odonata) from Costa Rica. — Tijdschrift voor Entomo-
logie 132: 163-176, figs. 1-24. [ISSN 0040-7496]. Published 1 December 1989.
Palaemnema baltodanoi
(Platystictidae),
Philogenia
peacocki (Megapodagrionidae),
Phyllogomphoides burgosi and Epigomphus echeverrii (Gomphidae) are described from a
collection of Odonata made in Guanacaste National Park, north-west Costa Rica. A fifth new
species, Epigomphus houghtoni from eastern Costa Rica, is also recorded. The possible affinities
of the new species are discussed.
Key words. — Odonata, Costa Rica, Guanacaste, new species.
Mr. Stephen J. Brooks, Department of Entomology, British Museum (Natural History),
Cromwell Road, London SW7 5BD
INTRODUCTION
This paper forms the first part of a larger project
to make an inventory of the Odonata of Guanacaste
National Park, in north-west Costa Rica, and ul-
timately to produce a field-guide to the dragonflies
of Costa Rica. Four of the five species described
below were part of a collection that I made in
Guanacaste National Park during July 1988. The
fitth species was collected by Mr. John Paul in
eastern Costa Rica and for convenience it is also
recorded here.
Most of the 700 km? of Guanacaste National
Park comprises tropical dry forest (Janzen 1986),
although all of the new species were collected in
the rain-forest covering the western slopes of two
volcanoes at the northern end of the Cordillera
de Guanacaste. Here there are many permanent
rivers and streams which support large assemb-
lages of Odonata. Some of these species are wide-
ranging throughout much of Mesoamerica but
some genera, and particularly those in which the
new species are described, include species which
have more restricted distributions. The odonate
fauna of Costa Rica is probably the best known
in all of Central America and comprises about 250
species (Paulson 1982). However, until I visited
Guanacaste National Park there had been no major
collection of Odonata on Volcan Orosi or Volcan
Cacao. Several of the species that I collected on
the volcanoes, such as Erpetogomphus tristani Cal-
vert, Perigomphus pallidistylus (Belle), Neocor-
163
dulia batesi longipolex Calvert and Argia rogersi
Calvert, had rarely been collected before. Of a total
of 15 species collected on Volcan Cacao, three were
undescribed. Therefore, it is possible that the new
species are endemic to the river systems on the
volcanoes or are at least restricted to the surviving
remnants of upland rain-forest in the northern
part of the country.
TAXONOMY
Palaemnema baltodanoi sp.n.
(figs. 1—5)
Material examined. — Holotype @: Costa Rica, Gua-
nacaste Province, SW side Volcan Cacao, Estacion Mengo,
1100 m, July 1988, S. J. Brooks. Paratypes 14 6, 3 2:
same data as holotype. Holotype and paratypes deposited
in BMNH.
Description of male holotype.
Head: width across eyes 4.7 mm. Eyes dark
brown in life. Labrum pale blue with black stripe
across anterior margin; base of mandible pale blue;
gena pale blue; anteclypeus pale blue; postclypeus
black; frons and area between eye and antenna
from lateral ocellus to gena metallic dark blue;
vertex black with small pale brown spot adjacent
to lateral ocellus; occiput black; labium pale brown.
Prothorax: anterior lobe apple green, black me-
dially; propleuron black; median lobe with large
apple green lateral spot; posterior lobe black.
164 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 1—5. Palaemnema baltodanoi sp.n. — 1, Pterothorax, left lateral view. 2, Male anal appendages and apex of
abdomen, dorsal view. 3, Male anal appendages and apex of abdomen, lateral view. 4, Penis, ventral view. 5, Female
apex of abdomen, lateral view.
BROOKS:
Pterothorax (fig. 1): bronze-black on dorsum
with narrow (width 10% of episternum 2) apple
green antehumeral stripe extending from collar
carina to within 10% of antealar carina; mesepi-
meron black in dorsal half, apple green below, this
pale area tapering anteriorly with broad black
stripe adjacent to antealar carina; metepisternum
apple green in dorsal half, black ventrally; metepi-
meron apple green, becoming pale yellow ven-
trally, with narrow black stripe adjacent to second
lateral suture.
Legs: white with black stripe on dorsum of
femur and fore tibia; coxa white; tarsus black.
Wings: hyaline with yellow-brown suffusion
along costal margin, darkest at apex. Venation
black. Pterostigma red-brown with narrow pale
border; 1.4 mm in fore wing, 1.7 mm in hind wing;
2-2.5 cells below pterostigma in fore wing, 1.75 in
hind wing. 25 postnodals in fore wing, 20-22 in
hind wing. R’ arises at 8th postnodal in fore wing,
7th in hind wing.
Measurements: hind wing 28.0 mm; 5.0 mm at
greatest width.
Abdomen: black with the following pale yellow
ventro-lateral markings: $1 with large square spot;
S2 with longitudinal stripe, tapering apically, in
basal three-quarters; S3-S7 with tapering longi-
tudinal stripe in basal quarter; S8 with rounded
spot in basal quarter; S9-S10 unmarked. Penis fil-
ament tips (fig. 4) form A of Calvert (1931).
Superior appendages (figs. 2-3): black; about
same length as inferiors, three times as long as S10;
parallel in basal half, converging apically with
short, blunt submedian (53% of length) tooth on
inner dorsal margin; appendage abruptly widened
ventrally at 70% of length with ventral margin of
this portion straight; apex straight.
Inferior appendages (figs. 2-3): black with white
ventro-lateral subapical spot; diverging basally but
strongly incurved in apical 20%, this latter section
with transverse ridges on inner surface; abruptly
constricted at apex to give short, curled apical
tooth; ventral margin with slight swelling present
just proximal to apical tooth; short, blunt, apically
projecting tooth present on inner margin at 25%
of length; dorsum of basal 25% concave; in lateral
view appendage angled dorsally at about 45°.
Measurements: total length 47.5 mm; abdomen
39.5 mm (including appendages 1.3 mm).
Description of female.
As male except the following:
Prothorax: pale markings dull yellow-green.
Pterothorax: pale markings dull yellow-green,
mesepimeron entirely black.
Dragonflies from Costa Rica 165
Wings: hind wing 26.0-27.5 mm. Pterostigma
1.4 mm in fore wing, 1.5 mm in hind wing; 25-27
postnodals in fore wing, 21-23 in hind wing.
Abdomen: pale markings yellow-brown, trian-
gular, broader and shorter than in male; S9 with
large lateral blue-grey spot in basal two-thirds ex-
tending dorsally in basal half. Ovipositor extending
0.4 mm beyond apex of abdomen; straight ven-
trally, untoothed (fig. 5).
Anal appendages: short, conical, 0.4 mm in
length.
Measurements: total length 40.5-44.0 mm; ab-
dominal length 32.5-36.0 mm.
Variation in male paratypes. — Total length
43.0-55.0 mm; hind wing 25.5-30.5 mm; pteros-
tigma 1.2-1.4 mm; fore wing postnodals 24-27.
Antehumeral stripe sometimes longer than in hol-
otype but never confluent with antealar carina.
Habits. — All specimens were collected over the
riffle reach of a shallow, narrow stream on a steep
incline in densely shaded rain-forest. Their dark
coloration made them difficult to locate in the
gloom. Specimens were numerous and were the
commonest odonate at the locality. Other species
present were Cora chirripa Calvert, Hetaerina ma-
juscula Selys, Philogenia peacocki sp.n.,
Epigomphus sp. indet. and Brechmorhoga rapax
Calvert.
The species was reluctant to fly and was usually
encountered hanging from the leaves and branches
of trees and bushes adjacent to the stream from
which they could be removed by hand. The flight
was weak, slow and of short duration. Two males
were seen to display to one another. A perched
male took off and confronted a second male which
had attempted to land on the same perch. Both
hovered at a height of about 1 m facing each other,
with the pale blue clypeal stripe clearly visible,
slowly moving backwards and forwards in unison
keeping about 0.3 m apart. After about one minute
the intruder retired and the first male returned to
the perch. Females were never seen in tandem with
males but were found perching nearby.
Remarks. — Paulson (1982) lists 17 species of
Palaemnema from Central America of which seven
are known from Costa Rica. There is a high degree
of endemism with six of the seven Costa Rican
species being endemics. Typically these species in-
habit shady streams in dense forest although the
more widely distributed species occur in more
open, lowland localities.
Males of Palaemnema baltodanoi sp.n. can be
166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 6-9 Philogenia peacocki sp.n., male holotype. — 6, Pterothorax right lateral view. 7, Anal appendages, ventral
view. 8, Apex of abdomen and anal appendages, interno-dorsal view. 9, Apex of abdomen and anal appendages, lateral
view.
BROOKS:
distinguished from most of the other species in the
genus by the lack of blue markings at the apex of
the abdomen. Also, the abrupt median swelling of
the ventral margin of the superior appendages is
distinctive since in most species this gradually wid-
ens. In the keys to the genus provided by Calvert
(1931) the new species comes out to apicalis Cal-
vert. However, unlike baltodanoi, there is no an-
tehumeral stripe in apzcalis and the medio-dorsal
tooth of the superior appendage is not as promi-
nent in lateral view. The apical tooth of the inferior
appendage in apicalis is long and abruptly curved
dorsally (like a hairpin) to form a narrow dorsal
notch but in baltodanoz it is shorter, more rounded
and curves inwards with no notch on the dorsal
margin. The inferior margin of the superior ap-
pendage, where it expands in the apical half, is not
angulate in apicalis and is narrower than in
baltodanoi. At the apex of the penis filament there
is a basal lobe in apicalis which is absent in
baltodanoi.
The anal appendages of baltodanoi are similar
morphologically to P pazlirica Calvert and
distadens Calvert and in particular there is a swel-
ling on the ventral margin of the inferior appen-
dage just proximal to the apical hook in both
distadens and baltodanoi. However, paulirica
differs from baltodanoi in possessing a slight in-
vagination at the apex of the superior appendages
and in distadens the apical tooth on the inferiors
is not curled. The thoracic markings of baltodanoi
are also similar to distadens but in distadens and
paulirica the abdominal segments 8 and 9 are
marked blue dorsally.
This species is named in honour of Sr Jorge
Baltodano who very kindly allowed his ranch to be
purchased by Guanacaste National Park.
Philogenia peacocki sp. n.
(figs. 6—9)
Material examined. — Holotype &: Costa Rica, Gua-
nacaste Province, SW side Volcan Cacao, Estacion Mengo,
1100 m, July 1988, S. J. Brooks. Paratype @: Costa Rica,
Puntarenas Province, Monteverde, Rio Guacimal and
small streams in biological reserve, 1600 m, 7 June 1986,
T. W. Donnelly. Holotype deposited in BMNH, paratype
in Donnelly collection.
Description of male holotype.
Head: width across eyes 6.7 mm. Eyes black in
life. Labrum pale blue; base of mandible pale blue;
gena pale blue; anteclypeus and postclypeus black;
frons black with small pale blue spot below scape;
vertex black with pale blue spot between lateral
ocellus and eye; labium and occiput black.
Dragonflies from Costa Rica 167
Prothorax: anterior lobe with blue medio-lateral
spot; median lobe with large blue lateral spot; pos-
terior lobe with small blue lateral spot.
Pterothorax (fig. 6): azure blue with the follow-
ing black markings: broad antehumeral stripe ex-
tending over dorsal carina; small spot at posterior
end of humeral suture; narrow black stripe adjacent
to antealar carina with short stripe extending onto
first and second lateral suture; katepisternum 2
black; broad black stripe at anterior end of mese-
pimeron; katepisternite 3 black. Minute blue spots
on antealar sinus. Notum of thorax with pale
blue/grey pruinescence.
Legs: femur blue with black stripe on posterior
side; tibia and tarsus black with pale brown stripe
on posterior side of mid and hind leg.
Wings: hayline with yellow-brown suffusion es-
pecially around margins. Venation black. Pteros-
tigma dark reddish brown, 2.2 mm in fore wing, 2.5
mm in hind wing. 24-25 postnodals in fore wing,
22-23 in hind wing. 4-5 cells below pterostigma;
R’ arises between 10th and 11th postnodal in fore
wing, between 8th and 9th in hind wing.
Measurements: hind wing 34 mm, greatest
width 6 mm.
Abdomen: black with the following markings:
S1 with broad lateral blue spot, S2-S7 with small
blue oval baso-lateral spot and yellowish tapering
stripe in basal quarter; blue-grey pruinosity on dor-
sum and laterally on S1-S2 and S9-S10.
Superior appendages (figs. 7-9): black with dor-
sal blue-grey pruinosity; strongly convergent in
dorsal view; apical half downcurved in lateral view;
meso-ventral process absent; apex swollen with
shallow median indentation.
Inferior appendages (figs. 7-9): 0.7 times length
of superiors; parallel not diverging; strongly curv-
ing dorsally in lateral view, tapering to blunt apex
with a small subapical notch; setose, dorsally pro-
jecting, basal tubercle present.
Measurements: total length 51 mm; abdomen
40.5 mm (including anal appendages 1.4 mm).
Female unknown.
Variation in paratype. — 25-26 postnodals in
fore wing; 23-24 postnodals in hind wing. Hind
wing 36 mm, 6.5 mm at widest. Total length 55
mm; abdomen 42 mm. When viewed laterally the
internal apical lobe of the superor appendages ex-
tends further ventrad. The superiors, on the in-
terno-dorso-lateral margin, have a more promi-
nent lobe just proximal of the ventral expansion.
The subapical tooth on the inferiors is smaller. In
dorsal view, the median dog-leg curve of the infe-
rior appendage is more angulated.
168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Habits. — The holotype was perched on the
branch of a bush at about 2 m, overhanging a riffle
reach on a narrow (1 m), shallow (0.2 m) stream
in densely shaded rain-forest. It was the only spec-
imen seen.
Remarks. — Six species of Philogenia are known
from Central America of which three occur in
Costa Rica (Bick & Bick 1988, May 1989). Only one
of these is endemic to the country but the other two
species occur only in Costa Rica and Panama. In
fact, as Bick & Bick (1988) point out, only one of
the 28 known species of Philogenia has anything
like a widespread distribution. The distribution of
the genus is centred on northern South American
with Costa Rica being its northern limit.
The new species belongs to the helena (Hagen)-
group of species (Bick & Bick 1988). It seems clos-
est to P. berenice Higgins, described from Peru, but
the superior appendages are relatively longer and
more abruptly downcurved in berenice and the
meso-ventral process is broader in the latter spe-
cies. The inferior appendages of berenice lack the
ventral cavity and dorso-basal tubercle, although
the dorso-basal ridge is more developed. In addi-
tion, at the apex of the inferior appendages of
berenice are a pair of small, subequal tubercles,
rather than a subapical notch which is present in
peacocki.
This species is named in honour of Mr. Harold
Peacock who very kindly allowed his ranch to be
purchased by Guanacaste National Park.
Phyllogomphoides burgosi sp.n.
(figs. 10—13)
Material examined. — Holotype 9: Costa Rica, Alaju-
ela Province, 8 km S Santa Cecilia, Estacion Pitilla, 680
m, July 1988, S. J. Brooks. Holotype deposited in BMNH.
Male: Unknown.
Description of female holotype.
Head (fig. 10): width across eyes 10.2 mm. Eyes
blue-grey in life. Labrum black with medio-lateral
yellow-green spot; mandible black, yellow-green at
base; gena brown with small yellow-green ventral
spot; anteclypeus yellow-green; postclypeus brown
with yellow-green lateral spot; frons brown with
prominent lateral, dorsally projecting horn 2.8 mm
in height, 1.5 mm wide at base, 0.4 mm at apex;
horn brown with pale yellow-green stripe on inner
and front surface and tuft of short, posteriorly
projecting setae at apex; vertex red-brown with
steep M-shaped ridge behind lateral ocelli; dorsum
of occiput dark brown with shallow median depres-
sion and fringe of long setae on upturned posterior
margin; occiput red-brown with shallow groove
extending from occipital tubercle to sinus of pre-
mandibular suture, crossed at right angles by nu-
merous short striae.
Prothorax: red-brown with yellow lateral spot
on anterior lobe, yellow median and lateral spot on
middle lobe.
Pterothorax (fig. 11): red-brown with the fol-
lowing yellow markings: narrow stripe on dorsal
carina; collar carina with broad stripe; antehumeral
stripe short and narrow; humeral stripe broaden-
ing posteriorly; broad mesepimeral stripe; metep-
isternal stripe broad, interrupted at spiracle with
isolated spot anterior of spiracle; katepisternite
with large spot; metepimeral stripe narrowly mar-
gined with brown; sternites with median stripe.
Legs: tibia and tarsus black; femur yellow-brown
but fore femur black on outer surface.
Wings: hyaline with amber suffusion around
veins, becoming darker at wing base. Venation
black. Pterostigma dark red-brown, 5.4 mm in fore
wing, 5.9 mm in hind wing. Basal subcostal cross-
vein present. Nodal index 17 : 25 : 23 : 16/15 : 18
: 18 : 17. Secondary primary antenodal 8th or 9th
crossvein in fore wing, 8th in hind wing. Inter-
median crossveins 14 : 13/11 : 10. Supratriangle
2-celled in fore wing, 2-3 celled in hind wing.
Discoidal triangle 3-celled; subtriangle 3-celled.
One cubitoanal crossvein in all wings. Anal loop
3-celled.
Measurements; hind wing 47 mm, greatest
width 12 mm.
Abdomen: black with the following yellow
markings: Sl dorsum with small medio-apical spot
and lateral spot in ventral half; S2 dorsum with
basal T-shaped marking extending to second car-
ina, laterally with spot in ventral half; S3 with
lateral semicircular spot almost extending to trans-
verse carina, dorsum marked with fine mid-dorsal
stripe broadening slightly at base; S4 marked dor-
sally with small basal spot and laterally with small
dome-shaped spot terminating before transverse
carina; SS-S6 marked similarly to S4 but spots
progressively smaller; S7 with basal band extend-
ing to transverse carina; S8 with wedge-shaped
medio-ventral lateral spot and minute isolated spot
beyond apex of wedge; S9-S10 unmarked. Folia-
tions absent. Vulvar scale (fig. 12) broadly U-
shaped, arms broad with small apical tubercle, 0.56
mm in length.
Anal appendages (fig. 13): black in basal 0.15%
then yellow, lanceolate with black tip.
Measurements: total length 70 mm; abdomen 51
mm (including anal appendages 3.1 mm).
BROOKS:
Dragonflies from Costa Rica 169
Figs. 10—13, Phyllogomphoides burgosi sp.n., female holotype. — 10, Head, lateral view. 11, Pterothorax, right lateral
view. 12, Vulvar scale, ventral view. 13, Apex of abdomen, lateral view.
Habits. — The specimen was collected flying in
rain-forest over a narrow slow-flowing, shaded
stream and was the only specimen seen.
Remarks. — A total of ten species of
Phyllogomphoides have been recorded from Cen-
tral America (Paulson 1982) of wich four are
known from Costa Rica. Most of the Central Amer-
can species are quite widespread and only three are
restricted to just one country. None of the pre-
viously described species is restricted to Costa Rica
but P. appendiculatus Kirby occurs only in Costa
Rica and Panama.
The new species appears to belong to the
bifasciatus (Hagen)-group (Donnelly 1979) in
which the vulvar scale is simple without lateral
spines or protrusions. It is apparently related to the
Trinidadian species P. cornutifrons (Needham)
which is the only other known Phyllogomphoides
species to have frontal horns in females. However,
in P. cornutifrons the frontal horns are much more
slender and elongate, and the branches of the vul-
var scale are more rounded apically than in the new
species. In addition P. burgosi is considerably larger
than cornutifrons. Although the male of burgost is
still unknown there can be little doubt concerning
the identity of the species because of the possession
of the extraordinary frontal horns. I feel that it is
justifiable to describe the species based on a single
female specimen in order to draw attention to the
existence of second Phyllogomphoides species
with frontal horns.
Of the Central American species only the female
of P. pugnifer Donnelly is unknown. However, this
170 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 14—17. Epigomphus echeverrii sp.n., male holotype. — 14, Pterothorax, right lateral view. 15, Apex of abdomen
and anal appendages, dorsal view. 16, Left anal appendages, interno-caudal view (right appendages not shown). 17,
Apex of abdomen and anal appendages, lateral view.
BROOKS:
species is considerably smaller than burgosi and the
male pterothoracic markings differ. In pugnifer the
antehumeral stripe is confluent with the stripe
near the collar carina and the humeral and mesep-
imeral stripes are narrower than in burgost.
This species is named in honour of Sr. Mario
Burgos who very kindly allowed his ranch to be
purchased by Guanacaste National Park.
Epigomphus echeverrii sp. n.
(figs. 14—21)
Material examined. — Holotype @: Costa Rica, Gua-
nacaste Province, SW side Volcan Cacao, Estacion Mengo,
1100 m, July 1988, S. J. Brooks. Paratypes 1 8, 2 9 data
same as holotype; 1 ® Costa Rica, Guanacaste Province,
W side of Volcan Orosi, Estacion Maritza, 600 m, July
1988, S. J. Brooks. All specimens deposited in BMNH.
Description of male holotype.
Head: width across eyes 8.1 mm. Eyes turquoise
in life. Labrum black; base of mandible yellow with
small brown basal spot; gena pale blue; anteclypeus
and postclypeus brown; frons brown anteriorly,
pale blue-green dorsally with narrow basal and
broad median brown stripe; vertex and dorsum of
occiput dark brown; labium and occiput pale blue-
green.
Prothorax: dark brown with small lateral, me-
dian and anterior blue-green spots on median lobe.
Pterothorax (fig. 14): black with the following
pale blue-green markings: stripe adjacent and pa-
rallel to collar carina; short, narrow antehumeral
stripe; two elongate humeral spots close to hu-
meral suture, one near anterior suture, the second
near the antealar carina; narrow mesepimeral
stripe; metepisternal stripe interrupted posteriorly
leaving small isolated spot near antealar carina;
metepimeron with broad stripe concurrent poste-
riorly with narrow ventral stripe.
Legs: tibia and tarsus black, femur dark brown,
paler on posterior edge and internal face of fore
femur yellow-green. Hind leg with spines on outer
row of tibia and first two tarsal segments short,
thick, rounded apically; apical spines on outer edge
of fore tibia elongate.
Wings: hyaline with yellowish tinge. Venation
black. Pterostigma dark red-brown, 3.5 mm in fore
wing, 4.0 mm in hind wing. Basal subcostal cross-
vein present. Nodal index 16 : 20 : 20 : 16/16 : 15
: 15 : 16. Second primary antenodal 8th crossvein
in fore wing, 7th in hind wing. Intermedian cross-
veins 7 : 8/4 : 5. Supratriangle one-celled. Discoidal
triangle free in left fore wing, crossed in other
wings; subtriangle free. 3-4 cubitoanal crossveins
Dragonflies from Costa Rica AL
in fore wing, 3 in hind wing. Hind wing with 5
paranal cells, 4 postanal cells, 4 rows of cells pos-
terior to Cu.
Measurements: hind wing 36 mm, greatest
width 9.5 mm.
Abdomen: black with the following pale blue-
green markings: S1 pale laterally; S2 with narrow
mid-dorsal stripe, pale laterally; S3 with narrow
mid-dorsal stripe and lateral spot in basal three-
quarters; S4-S6 with narrow dorsal band at base of
each segment, and lateral spot tapering to trans-
verse carina; S7 entirely pale in basal two-thirds;
S8-S9 unmarked; S10 pale brown in basal half,
darker apically with tapering ventral spot.
Anal appendages (figs. 15-17): black; superior
appendages triangular in cross-section, about three
times as long as broad when viewed dorsally, 1.8
times as long as broad when viewed laterally, bif-
urcate apically, outer branch bending laterally tap-
ering to narrow point, inner branch curving ven-
trally with blunt apex; inferior appendage 0.6 mm
longer than superiors, deeply bifurcate with
branches divergent at base, parallel in apical half,
apices of branches with a pair of short upturned
teeth, base of branches with transverse ridge, ter-
minating in low dorsal projection at each end, outer
projection elongate, inner projection short, broad
with numerous apical setae.
Measurements: total length 58.4 mm; abdomen
43.8 mm (including anal appendages 2.4 mm).
Description of female.
As male except: head (figs. 18-19), width across
eyes 9.4 mm. Labrum dark brown with black ante-
rior margin; vertex with pair of large ovate plates
projecting beyond posterior margin of head; occi-
put with sinuous transverse groove extending from
postocciput to sinus in premandibular suture, this
groove crossed at right angles by several short,
shallow striae and about halfway along groove is
low, rounded projection (fig. 19).
Pterothorax: pale lateral stripes yellower than
male.
Legs: femur almost entirely yellow-brown. Setae
on tibia and tarsus of hind legs unmodified.
Wings: hyaline with yellowish suffusion at base
and in antenodal area. Pterostigma 4.4 mm in fore
wing, 4.7 mm in hind wing. Nodal index 13 : 20 :
19 : 15/14 : 15 : 15 : 13. Second primary antenodal
crossvein seventh crossvein in all wings. Interme-
dian crossveins 6 : 6/4 : 4. Discoidal triangle free.
All wings with 3 cubitoanal crossveins. Hind wing
with 3 postanal cells.
Measurements: hind wing 40.5 mm, 10.5 mm at
greatest width.
172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 18—21. Epigomphus echeverr sp.n., female. — 18, Head, dorsal view. Note puncture marks near inner margin
of eyes. 19, Right eye, rear view. 20, Vulvar scale, ventral view. 21, Apex of abdomen, lateral view.
BROOKS:
Abdomen: black with yellow-green markings; S3
with lateral pale marking narrowing beyond trans-
verse carina; S7 pale only in basal third; S10 en-
tirely black; abdomen with greyish pruinesence
ventrally. Vulvar scale (fig. 20) narrowly V-shaped
with narrow arms, 0.78 mm in length. Dried ova
ovate, 0.52 X 0.3 mm.
Anal appendages (fig. 21) lanceolate.
Measurements: total length 62 mm; abdomen 41
mm (including anal appendages 1.4 mm).
Variation in paratypes. — Labrum with postero-
lateral blue-green spot and antero-lateral lobes of
postclypeus black in male. Wings with distinct
amber suffusion in male and females, possibly
older specimens.
Habits. — I collected two males and one female
near Estacion Mengo by the riffle reach of a shallow
stream in semi-shaded rain-forest with a broken
canopy. They were flying in the company of
Epigomphus subobtusus Selys and were quite com-
mon at the locality while the sun was shining but
disappeared when it became overcast. In flight the
males characteristically curved the apical segments
of the abdomen ventrally. Like other Epigomphus
species, the males frequently perched on low horiz-
ontal sticks at the edge of the stream and flew close
to the surface of the water. The female was seen
flying low over the stream in a pool of sunlight.
The other Mengo female was collected at the side
of a track ina field cleared of trees about 2 km from
the edge of the forest. The insect was perched in
long grass, probably sheltering from the wind
which was blowing very strongly at the time, and
may have been blown there from the forest. The
Maritza female was discovered dead in the web of
an orb-web spider next to a riffle reach on a large
river in forest with broken canopy.
In all three females there was damage to the
inner dorsal margin of the eyes (fig. 18) which was
probably inflicted by the apical spines on the infe-
rior appendages of males during mating. These
mating marks or ocular cicatrices are usually super-
ficial in Odonata (Dunkle 1979) but more exten-
sive damage has been described in species of
Ophiogomphus (Dunkle 1984) and in three species
of Epigomphus (Calvert 1920). In the three fe-
males of E. echeverrii the ocular cicatrices took the
form of at least two rounded holes which had some-
times coalesced to form a longitudinal groove. The
spines at the apex of the inferior appendage in
males of echeverrii are positioned transversely
about 0.5 mm apart but the holes in the eyes are
situated one above the other about 0.2 mm apart.
Dragonflies from Costa Rica 173
This suggests that the hole in the eye was caused
by only the outer spine of the inferior appendage
and that the inner spine fits into the depression in
the vertex lobe between the lateral ocellus and the
outer lateral margin of the lobe. In one female
specimen a hole had been punctured in the vertex
lobe in this position. The presence of more than
one hole in each eye implies that each female had
mated at least twice.
Remarks. — Twelve species of Epigomphus
have been recorded from Central America (Paul-
son 1982, Donnelly 1986) of which seven occur in
Costa Rica, five as endemics. Most of the species
have a limited distribution and are confined to
forested streams.
Although none of the female specimens was
collected zn copula with a male, it is very likely that
they are conspecific since the pterothoracic mark-
ings of both sexes are very similar. Also, one of the
females was coliected flying on the same stretch of
stream at the two males where the only other
Epigomphus species was subobtusus. Females of
subobtusus do not possess the large plates on the
vertex which are characteristic of echeverri. Addi-
tionally, the females here described do not resem-
ble the females of E. armatus Ris or E. tumefactus
Calvert which were the only other species of
Epigomphus collected during my stay in Guana-
caste National Park. The females are unknown in
E. clavatus Belle, E. pechumani Belle and E. paul-
soni Belle but none of these species has been re-
corded from Costa Rica and the pterothoracic
markings are different from echeverru.
Epigomphus echeverrii sp.n. is probably closely
related to the crepidus-pechumani-subsimilis
group of species, all of which have broad superior
appendages which bifurcate apically and long, nar-
row, deeply bifurcate inferiors which project
beyond the apex of the superiors and bear a pair of
short teeth at the apex of each branch. However,
in all the latter species, unlike echeverri, one of the
teeth at the apex of the branches of the inferior
appendage is positioned slightly subapically. E.
echeverrii differs from subsimilis Calvert in which
the branches of the inferior appendages are rela-
tively short and the outer apical projection of the
superior appendage curls ventrally. E. pechumani
Belle has a basal internal spine on the superiors,
which is absent in echeverri, and the inner apical
fork is toothed. In addition, the inferiors are shor-
ter than those of echeverrii. In crepidus Kennedy
the outer apical branch of the superior appendage
is very short and blunt and, although the inferiors
are morphologically similar to echeverri, the inner
174 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 22-24. Epigomphus boughtoni sp.n., male holotype. — 22, Pterothorax, rigth lateral view. 23, Apex of abdomen
and anal appendages, lateral view. 24, Inferior appendage, dorsal view.
tooth at the apex of the branch of the inferiors is
about twice the length of the outer and the apex is
more acutely pointed. Although some specimens of
crepidus have two antehumeral stripes, in other
individuals the lower stripe is interrupted medially
as it is in echeverrit.
The females of E. echeverrii are easily distin-
guished from other species in the genus by the
large flattened lobes on the vertex which project
considerably from the rear of the head. There are
no similar structures in any of the species of the
genus in which the females are known. In females
of crepidus and subsimilis (the female of
pechumani is unknown) there is a longitudinal
groove adjacent to the lateral ocellus and a pair of
small tubercles on the vertex.
This species is named in honour of Sr Gustavo
Echeverri who very kindly donated his ranches to
the Guanacaste National Park.
Epigomphus houghtoni sp.n.
(figs. 22—24)
Material examined. — Holotype: 4, Costa Rica,
Limon, Siquirres, 5.iv.1988. J. Paul. Holotype deposited in
BMNH.
Description of male holotype.
Head: width across eyes 7.8 mm. Colour of eyes
BROOKS:
in life unknown (probably blue). Labrum black
with postero-lateral yellow-green spot and small
pale brown median spot; base of mandibles yellow-
green; genae yellow-green; postclypeus dark
brown, marked black medially and on antero-lat-
eral lobes; frons dark brown anteriorly, pale yel-
low-green dorsally with basal and median black
stripe; vertex and occiput black; labium and back of
eyes yellow-green.
Prothorax: black with median and lateral pale
green spot on middle lobe.
Pterothorax (fig. 22): black dorsally, dark brown
laterally; marked dorsally with pale green trans-
verse stripe adjacent to collar carina, short pale
green antehumeral stripe and pale green posterior
spot positioned between antehumeral stripe and
humeral suture; marked laterally with narrow yel-
low mesepimeral stripe, yellow metepisternal
stripe broad anteriorly narrowing posteriorly,
metepimeron almost entirely yellow with narrow
brown stripe adjacent to posterior half of second
lateral suture and ventral brown stripe in anterior
half.
Legs: almost entirely dark brown-black with pale
brown stripe on posterior side of femur. Hind leg
with spines on outer row of tibia and first two
tarsal segments short, thick, rounded apically.
Wings: membrane hyaline with slight yellow-
brown suffusion at base of wings and along poste-
rior margin of hind wing. Venation black. Ptero-
stigma dark reddish brown, 3.7 mm in fore wing,
4.2 mm in hind wing. Basal subcostal crossvein
present. Nodal index 15 : 19: 18: 14/14: 13 : 14
: 14. Second primary antenodal crossvein seventh
in fore wing and hind wing. Intermedian cross-
veins 5-6/3-3. Supratriangle one celled. Discoidal
and sub-triangle uncrossed. 3-4 cubitoanal cross-
veins in fore wing, 2 in hind wing. Hind wing with
5 paranal cells, 3 postanal cells, 3 rows of cells
posterior of Cu.
Measurements: hind wing 34.5 mm, greatest
width 9.5 mm.
Abdomen: dark brown with the following yellow
markings: Sl and S2 with narrow medio-dorsal
stripe and entirely yellow laterally; S3 with narrow
median dorsal stripe restricted to basal half, large
lateral spot in lower half occupying basal three-
quarters of segment; Sá-S6 with lateral spot in
lower half of segment extending to lateral carina
and small mid-dorsal basal spot; S7 entirely yellow
in basal two-thirds; S8-S9 with small spot on baso-
lateral membrane; S10 marked with tapering ven-
tral spot.
Anal appendages (fig. 23-24): black; superior
appendages rectangular, about 2% times as long as
Dragonflies from Costa Rica 175
broad, dorsal edge cuving ventrally in apical half
with seven short teeth on ventro-apical margin;
inferior appendage with dorso-basal depression;
deeply bifurcate; each branch with low medio-dor-
sal ridge and small median spine in shallow depres-
sion; apices hooked and curving inwards.
Measurements: total length 53.6 mm; abdomen
39.5 mm (including anal appendages 2.7 mm)
Remarks. — E. houghtoni belongs to the largest
species group in Epigomphus all of which have the
superior appendages rounded apically, bearing a
row of short subapical teeth or crenellations. Most
of these species, such as armatus Ris and
subobtusus Selys, have two antehumeral stripes but
a few, like occipitalis Belle and tumefactus Calvert,
have only one. Of this latter group houghtoni
seems to be closest to occipitalis since this is the
only other species in the genus which has a horse-
shoe-shaped inferior appendage. However,
occipitalis (which was described from Peru) can be
readily distinguished from houghtoni by the pres-
ence of a strong median tooth on each branch of the
inferior appendage which is absent in houghtont.
This species is named after Mr Greg Houghton
who accompanied John Paul in Costa Rica but who
was tragically killed shortly afterwards.
ACKNOWLEDGEMENTS
I am indebted to Dr. D. H. Janzen for assistance
and hospitality during my stay at Guanacaste Na-
tional Park. I am grateful to Drs. J. Belle (Velp, The
Netherlands), T. W. Donnelly (Binghamton Uni-
versity, USA) and R. Garrison (Azusa, USA) for
their helpful advice and ready exchange of infor-
mation. Thanks are also due to Dr. D. Azuma
(Academy of Natural Sciences, Philadelphia,
USA), Dr. T. W. Donnely, Mr. M. F. O'Brien (Uni-
versity of Michigan, Ann Arbor, USA), Mr. J. Paul
(Oxford, UK), Mr. S. R. Shaw (Museum of Com-
parative Zoology, Cambridge, USA) and Mr. J. van
Tol (Rijksmuseum van Natuurlijke Historie,
Leiden, The Netherlands) for the loan of speci-
mens.
REFERENCES
Bick, G. H. and J. C. Bick, 1988. A review of the males of
the genus Philogenia, with descriptions of five new
species from South America (Zygoptera: Megapoda-
grionidae). — Odonatologica 17 (1): 9-32.
Calvert, P. P, 1920. The Costa Rican species of
Epigomphus and their mutual mating adaptations. —
Transactions of the American Entomological Society
46: 323-354.
176 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Calvert, P. P, 1931. The generic characters and the species
of Palaemnema (Odonata: Agrionidae). — Transac-
tions of the American Entomological Society 57: 1-
TI
Donnelly, T. W., 1979. The genus Phyllogomphoides in
Middle America (Anisoptera: Gomphidae). — Odo-
natologica 8 (4): 245-265.
Donnelly, T. W., 1986. Epigomphus westfalli spec. nov.,
a new dragonfly from Nicaragua. — Odonatologia 15
(COTE UE
Dunkle, S. W., 1979. Ocular mating marks in female
nearctic Aeshnidae (Anisoptera). — Odonatologica 8
(2): 123-127.
Dunkle, S. W., 1984. Head damage due to mating in
Ophiogomphus dragonflies (Anisoptera: Gomphi-
dae). — Notulae Odonatologicae 2 (4): 63-64.
Janzen, D. H., 1986. Guanacaste National Park: tropical
ecological and cultural restoration. — Editorial Uni-
versidad Estatal a Distancia, San José, Costa Rica.
May, M. L., 1989. Status of Philogenia leonora Westfall
& Cumming (Zygoptera: Megapodagrionidae). —
Odonatologica 18(1): 95-97.
Paulson, D. R., 1982. In: Hurlbert, S. H. and A. Villalo-
bos-Figueroa [eds]. Aquatic biota of Mexico, Central
America and the West Indes, Odonata. pp. 249-277. —
San Diego State University, San Diego, California,
USA.
Received: 6 July 1989
Accepted: 8 July 1989
GENETIC DISTANCE IN THE GENUS EPHIPPIGER
(ORTHOPTERA, TETTIGONIOIDEA) - A RECONNAISSANCE
by
L. OUDMAN, W. LANDMAN & M. DUIJM
ABSTRACT
Oudman, L., W. Landman & M. Duijm, 1989. Genetic distance in the genus Ephippiger
(Orthoptera, Tettigonioidea). — a reconnaissance. — Tijdschrift voor Entomologie 132: 177-
181, figs 1-2, tabs 1-3. [ISSN 0040-7496]. Published 1 December 1989.
Genetic distances were determined by means of enzyme electrophoresis for a number of
Ephippiger (sub)species, mainly from southern France and northern Italy. For each (sub)species
and form one ‘typical’ location was selected. The results are summarized in a dendrogram.
The three groups distinguished by Duijm & Oudman (1983) on the base of copulatory behaviour
and morphological characters are confirmed. Nei's genetic distances between E. ephippiger,
E. cruciger and E. cunti appeared to be low for genuine species. A comparison with Uromenus
rugosicollis is included.
Keywords. - Ephippiger, enzyme electrophoresis, genetic distance, dendrogram.
L. Oudman, Department of Genetics, University of Groningen, P.O. Box 14, 9750 AA Haren,
The Netherlands; W. Landman, Noorder Dierenpark/Zoo, Emmen, The Netherlands; M.
Dutjm, Department of Zoology, University of Groningen; present adress: Onnerweg 41, 9751
VB Haren, The Netherlands.
INTRODUCTION
In France, the northern part of Spain and NW
Italy a number of Ephippiger (sub)species occur
(e.g. Chopard 1951; Harz 1969). For this region
Duijm & Oudman (1983) recognised three groups
of (sub)spieces, viz. 1 - Ephippiger provincialis
(Yersin, 1854), 2 - E. terrestris (Yersin, 1854) with
the three (sub)species E.t. terrestris, E. t. bormansi
(Brunner von Wattenwyl, 1882) and E. #. caprai
Nadig, 1980, 3 - E. ephippiger (Fieber, 1853) (the
subspieces E. e. vitium (Serville, 1831)!) and E.
e. vicheti Harz, 1966) E. cunti (Bolivar, 1877) and
E. cruciger (Fieber, 1853). Between these groups
no mating is possible. Within group 2 no mating
barriers were found, whereas mating between spe-
cies of group 3 is possible (Hartley & Warne 1984),
but not in all cases (Duijm & Oudman 1983). Iden-
tification of single specimens and even populations
belonging to group 3 often meets with considerable
difficulties owing to the large variability of the
morphological characters. The present study is an
attempt to elucidate the relations between these
Ephippiger taxa by the investigation of enzyme
polymorphism. For comparison the Ephippigerid
Uromenus rugosicollis (Serville, 1839) is used.
177
MATERIALS AND METHODS
The insects were collected during field trips in
August and September of 1979, 1980, 1981, 1982
and 1983. Generally we succeeded in collecting a
sufficient number (c 20) from a restricted area
of a few acres. This area had to be small to limit
ourselves to one population (or part of it) and
so to avoid the mixing of different populations.
The animals were killed, measured, photographed
and frozen in solid carbon dioxide (-79°C).
For this “reconnaissance” it appeared desirable
to omit the intra- (sub)specifical variation in order
to geta clearer picture. For each taxon we therefore
selected one locality that we considered sufficiently
typical. In this selection we used - if possible —
the type locality or our nearest collecting site and
otherwise a locality that was in good concordance
with the morphological description and/or in the
neighbourhood of the centre of distribution’). The
') According to Kruseman (1988) the correct name for
this subspecies is E. ephippiger diurnus Dufour, 1841.
2) Later work (Landman et al. 1989) showed that the
population used in this study as representative for
E. t. terrestris (Col de Castillon, No. 2), though in
many respects very close to the nominate form cannot
be regarded as entirely “pure”.
178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Fig. 1. Collection sites of Ephippiger and Uromenus
species in SW Europe. Numbers denote the sites, see
table 1.
sites selected are summarised in table 1 and fig. 1.
For preparation of the samples, electrophoretic
techniques and the preparation of the horizontal
polyacrylamide gels we refer to Van Dijk & Van
Delden (1981).
The following loci were analysed: Alcohol de-
hydrogenase (Adh), Tetrazolium oxidase (To), two
Phosphoglucomutases (Pgm-2 and Pgm-3), Malic
enzyme (Me), Fructosediphosphate aldolase (Ald),
Esterase-2 (Est-2), Hexokinase-3 (Hk-3), Fumar-
ate hydratase (Fum), Xanthine dehydrogenase
(Xdh), Isocitrate dehydrogenase (Idh), Glucose ox-
idase-3 (Gluo-3), a-Glycerophosphate dehydroge-
nase (a-Gpdh). For polymorphic loci Mendelian
inheritance was assumed on base of enzyme band
patterns and checked by testing genotype frequen-
cies for every location for Hardy-Weinberg equil-
ibrium.
Between populations Nei's genetic distances
were calculated (Nei 1975). From the matrix of
Table 1. Sites of collection of the selected populations of Ephippiger and Uromenus species.
No. (Sub)species Location
1. E. provincialis Plan d'Aups
2. Et. terrestris Col de Castillon
3. Et. terr. f minor Col de Maure
4. Et. caprat Cle. Scravaion
5. Et bormansı Naggio (L. di Como)
6. Ee. diurnus Cap Frehel
7. Ee. vicheti Naggio (L. di Como)
8. E. cruciger Gignac
9. E. cunti Cerbère
10. E. cunti f. jugicola Val d'Eyne
11. Uromenus rugosicollis Canigou
Table 2. Allele frequencies of the polymorphic loci of Ephippiger and Uromenus species.
(sub)species n
12 14
1 E. provincialis 16 0 0
2 Et. terrestris 18 50 50
3 Et.t.f. minor 13 25) 77
4 Et. caprai 23 .28 .65
5 Et. bormansi 18 .69 Bil
6 Ee. diurnus 20 0 0
7 Ee. vicheti DIL 0 0
8 E. cruciger 26 0 0
9 E. cuni 20 0 0
10 E. cunii f. jugicola 24 0 0
11 Uromenus rugosicollis
Pgm-2
Country/dept. Alt. m. Date
F 84 680 24-viii-80
F 06 700 31-viii-81
F 04 1350 24-vili-81
I Liguria 820 3-1x-81
I Lombardia 800 5/6-ix-81
1922 70 9-ix-83
I Lombardia 750 5/6-ix-81
F 34 60 3-vii-82
F 66 10 9-vii-82
F 66 1600 22-viii-82
F 66 840 31-viii-79
Pgm-3
16 20 26 29 32 35
0 1.00 Om 14 .86 0
0 0 U .67 0 .07
0 0 0 0 69 SI
‚07 0 0 .96 04 0
0 0 0 0 1.00 0
0 1.00 70 30 0 0
0 1.00 1.00 0 0 0
0 1.00 2» 23 31 oll il
0 1.00 .20 .80 0 0
0 1.00 0 1.00 0 0
- - 13 53 33 0
OUDMAN, LANDMAN & DUIJM:
genetic distances a dendrogram was constructed
following the UPGMA method (Sneath and Sokal
1976);
Samples of collected (sub)spieces from all local-
ities, including tips of abdomens used for electro-
phoresis, will be deposited in the Entomological
collection of the Institute for Taxonomical Zoology
(Zoological Museum) in Amsterdam.
RESULTS
Thirteen loci were investigated of which eight
were monomorphic. Five loci showed polymor-
phism: Est-2, Pgm-2, Pgm-3, To and Adh, with
respectively 4, 4, 4, 3 and 2 alleles. The allozyme
frequencies are given in table 2. To is fixed in most
populations. Adh is only polymorphic in E. #. ca-
prat. Pgm-2 is only variable in E. terrestris. The
most variable enzymes are Pgm-3 and Est-2.
The genetic distances are shown in table 3 and
the dendrogram, calculated from these distances, in
fig. 2.
Based on a preliminary investigation (Landman,
1981) we determined the genetic distance between
Uromenus rugosicollis and a number of Ephip-
piger (sub)spieces (18 populations, 9 loci, 18 al-
leles) at 0.3473. An indication of this distance is
added to the dendrogram.
DISCUSSION
The genetic distance (0.35) between the closely
related genera Uromenus and Ephippiger appears
to be very low in view of the range for genera
mentioned in reviews (e. g. Thorpe 1982, Menken
& Ulenberg 1987). The distance between genera
generally is 1, in the mean 1.30, and minimally
0.62.
The distance found by us between E. provincialis
and the other Ephippiger species (0.20) as well as
(Table 2 continued)
Genetic distance in Ephippiger 179
the distance between our groups 2 and 3 (0.16) are
very low for congeneric species. According to
Thorpe (1982) only in 3% of the cases studied
distances below 0.16 are found for congeneric spe-
cies.
The distances between the subspecies of E. ter-
restris (0.015 —0.11) are within the range generally
found between subspecies: 0.02 — 0.22 (Menken &
Ulenberg 1987) except one: the distance (0.015)
between Et. caprai and E.t. terrestris from Col de
Castillon. This very small distance is one of the
indications that the population of Castillon is not
quite representative for the nominate form of E. t.
terrestris.
The distances between the species within our
group 3 are strikingly small (0.03); they lie in the
range for subspecies. This is in accordance with the
results of Hartley and Warne (1984).
The dendrogram of fig. 2 offers a picture that is
mainly in accordance with current taxonomical
opinion. It also conforms to our grouping (Duijm
& Oudman 1983) based on morphological data and
on the existence of mating barriers. Within group
3, however, E. e. vicheti occupies a rather separate
position '). The relations between E. e. diurnus, E.
cruciger and E. cunit as well as those within the
terrestris-group will be dealt with in later publica-
tions.
ACKNOWLEDGEMENTS
We would like to thank the Uyttenboogaart-
Eliasen Stichting in Amsterdam for financial sup-
port. We are particularly indebted to Marten Zijl-
stra and Klaas Vrieling who helped us with the
') Nadig (1987,p.331) raises this taxon to species-level:
E. vicheti Harz, 1966.
Est-2 To Adh
15 17 20 23 40 63 65 67 18 22
.78 .22 0 0 0 .00 0 0 1.00 0
.08 .67 25 0 0 0 0 1.00 1.00 0
‚42 23 25 0 0 0 0 1.00 1.00 0
.10 DI 229 0 0 0 0 1.00 .80 .20
0 .24 .76 0 0 0 0 1.00 1.00 0
0 1.00 0 0 0 0 0 1.00 1.00 0
0 0 ‚08 192 0 0 0 1.00 1.00 0
.04 50 ‚46 0 .06 0 0 94 1.00 0
25 .67 .08 0 0 0 0 1.00 1.00 0
.08 55) 54 ‚04 0 0 0 1.00 1.00 0
0 .02 .90 ‚08 0 0 1.00 0 1.00 0
180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
genetic distance
0.4 0.2
(0)
SS o oe SSS _oc-
-crccsctobecttiteston
computing work. We thank Prof. W. van Delden
for providing laboratory facilities and for valuable
discussions and continued interest. Our thanks are
due to H. van Dijk for his instruction in electropho-
retic techniques. We are very grateful to Dr. A.
Nadig (Chur), who generously supplied us with
information about his collecting sites. We thank A.
Decae, G. J. van Klinken, H. Hidding and G. Over-
kamp, who assisted in collecting the insects. We are
grateful to Dr. M. G. Ritchie for critically reading
the manuscript and for correcting the English.
Furthermore we thank Mrs. J. Poelstra for typing
the manuscript and D. Visser for making the draw-
ings.
GROUP
Uromenus rugosicollis
Canigou
Ephippiger provincialis — 1
Plan d'Aups
Ephippiger eph. vicheti
Naggio
Ephippiger cunii
Cerbère
Ephippiger cunii f.jugicola | 3
Val d'Eyne
Ephippiger cruciger
Gignac
Ephippiger eph. diurnus
Cap Frèhel |
Ephippiger terrestris caprai
Scravaion
Ephippiger t. terrestris
Col de Castillon
3 5 F2
E.t. terrestris f. minor
Col de Maure
E.t.bormansi
Naggio. |
Fig. 2. Dendrogram of Ephippige-
ridae based on the genetic distances
of table 3. An estimation of the dis-
tance between Uromenus and the
Ephippiger (sub)species is added
on the base of preliminary work
(Landman, not published).
Table 3. Genetic distances based on ailozyme frequencies of thirteen loci of Ephippiger species.
Group (sub)species 1 2 3
1
2
E. provincialis = 2522 .1840
E.t. terrestris = .0680
E.t.t. f. minor =
Et. caprat
E.t. bormansi
Ee. diurnus
E.e. vicheti
E. cruciger
E. cunu
E. cunti f. jugicola
ID
i
O © JA AN
4
OUDMAN, LANDMAN & DUIJM:
REFERENCES
Chopard, L., 1951. Orthopteroides. - Faune de France 56:
1-359.
Van Dijk, H. & W. van Delden, 1981. Genetic variability
in Plantago species in relation to their ecology. I.
Genetic analysis of the allozyme variation in P. major
subspecies. — Theoretical and Applied Genetics 60:
285-290.
Duijm, M. & L. Oudman, 1983. Interspecific mating in
Ephippiger (Orthoptera, Tettigonioidea). - Tijdsch-
rift voor Entomologie 126: 97-108.
Ferguson, A., 1980. Biochemical Systematics and Evolu-
tion. — Blackie, London.
Hartley, J. C. & A. C. Warne, 1984. Taxonomy of the
Ephippiger ephippiger complex (ephippiger, cruciger
and cunit) with special reference to the mechanics of
copulation. - Eos 60: 43-54.
Harz, K., 1969. Die Orthopteren Europas, 1. - Series
Entomologica 5: i-xx, 1-749.
Kruseman, G., 1988. Matériaux pour la faunistique des
Orthoptères de France, Fascicule I, Les Ensiferes. —
Verslagen en Technische gegevens 51. Instituut voor
Taxonomische Zoölogie (Zoölogisch Museum), Uni-
versiteit van Amsterdam.
Landman, W. L., 1981. Genetische afstand en verwant-
schap in de Ephippigeridae. - Internal report Depart-
ment of Genetics, University of Groningen.
[unpublished].
Genetic distance in Ephippiger 181
Landman, W. L., L. Oudman & M. Duijm, 1989. Allozymic
and morphological variation in Ephippiger terrestris
(Yersin) 1854. — Tijdschrift voor Entomologie 132:
183-198.
Menken, S. B. J. & S. A. Ulenberg, 1987. Biochemical
Characters in Agricultural Entomology. - Agricultural
Zoology Reviews 2: 305-360.
Nadig, A., 1987. Saltatoria (Insecta) der Süd und Südos-
tabdachung der Alpen zwischen der Provence im W,
dem pannonischen Raum im NE und Istrien im SE.
I Tettigoniidae. - Revue suisse de Zoologie 94: 257-
356.
Nei, M., 1975. Molecular population genetics and evolu-
tion. - North Holland Publ. Co., Amsterdam.
Sneath, P. H. A. & R. R. Sokal, 1973. Numerical Taxon-
omy. — Freeman, San Francisco.
Thorpe, J. P., 1982. The Molecular Clock Hypothesis:
Biochemical Evolution, Genetic Differentiation and
Systematics. - Annual Review of Ecology and Syste-
matics 13: 139-168.
Received: 4 January 1989
Revised version accepted: 29 May 1989
182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
BOOK ANNOUNCEMENTS AND REVIEWS
Böcher, J., 1988. The Coleoptera of Greenland. - Med-
delelser om Gronland, Bioscience 26: 1-100, figs.
1-55. [ISBN 87-17-05572-5]. E. J. Brill, Leiden.
During a time that many entomologists focus
on the warmer parts of the world under the banner
of Biodiversity, it is good to see that the study
of polar regions on both sides of the globe is far
from neglected — and rightly so. This is undoubt-
edly due in part to the increasing accessibility of
these regions and to the increasing recognition
of their global ecological indicator value.
In the respected Danish series on Greenlandic
scientific research, Meddelelser om Groenland, an
interesting survey of the beetles of this largest
island of the world has recently appeared. It in-
cludes an extensively annotated, illustrated list of
the species (33 indigenous), with detailed biogeo-
graphical and ecological data and discussion. This
is all supported by maps, habitat descriptions, and
numerous references. Most of the species are qual-
ified as Lateglacial invaders from northern and
western Europe — probably passive invaders, as
several are flightless, but this view may be some-
what controversial. Genuine nearctic elements are
rare (one byrrhid species only). Most of the species
are distributed along the southern coasts. Of course
there are also synanthropic species and other in-
troductions; in fact about half of the total of nearly
70 is introduced by man, and includes such familiar
pests as species of Anthrenus, Attagenus, Ptinus,
Tribolium and Oryzaephilus.
The paleo-ecological implications of the recent
and fossil Greenlandic and other arctic beetle fau-
nas are amply discussed and compared with the
results of Coope and other workers on the Qua-
ternary history of the European, Siberian and
North American faunas. Judged from “in press”
references further reports on Plio-Pleistocene bee-
tle fossils are to be expected. Extreme environ-
ments like those of Greenland provide valuable
information on the tolerance of species to various
ecological factors, and highlight their suitability
as ecological indicators elsewhere in their range.
All in all, this survey of the Greenlandic Coleo-
ptera will prove to be extremely useful to ecol-
ogists, biogeographers, or the increasing number
of arctic naturalist travellers.
[J. Krikken]
Wirth, W. W. & W. L. Grogan, Jr., 1988. The predaceous
midges of the world (Diptera: Ceratopogonidae; Tribe
Ceratopogonini). — Flora and Fauna Handbook 4:
i-xvi + 1-160, figs. 1-49. [ISBN 0-916846-43-1].
E. J. Brill, Leiden. Price Hfl. 58.00.
The diagnostic characters of the genera pre-
viously assigned to the tribes Ceratopogonini and
Stilobezziini are reviewed and found not to support
a division into separate tribes. Forty-five genera
are combined into one tribe, the Ceratopogonini,
for which a diagnosis and a key to genera are
given. A diagnosis, illustrations, synonymy, and
a list of species are given for each genus and sub-
genus, but for the larger genera, species lists are
restricted to changes since the last published ca-
talogues or check lists.
[From the abstract by the authors].
ALLOZYMIC AND MORPHOLOGICAL VARIATION IN
EPHIPPIGER TERRESTRIS (YERSIN, 1854) (INSECTA,
ORTHOPTERA, TETTIGONIOIDEA)
by
W.LANDMAN, L. OUDMAN & M. DUIJM
ABSTRACT
Landman, W., L. Oudman, & M. Duijm, 1989. Allozymic and morphological variation in
Ephippiger terrestris (Yersin, 1854) (Insecta, Orthoptera, Tettigonioidea). — Tijdschrift voor
Entomologie 132: 183-198, figs. 1-13, tabs. 1-11. [ISSN 0040-7496]. Published 1 December
1989.
Based on allozyme frequencies the genetic distances between seventeen populations of Ephip-
piger terrestris are determined and compared with biometrics of epiproct, cerci and titillators.
The distinction of three subspecies, viz. terrestris, bormansi and caprai is confirmed. As a
result of the large variability of all characters within populations, distinction between subspecies
generally is not sharp. At the boundaries of the species distribution area the subspecies characters
are clearest, whereas in the centre an extensive area with intermediate populations is present,
generally exhibiting clinal differences. Such gradients are described for several characters for
a number of populations along a transect near Tende in the French Alps.
Key words. - Ephippiger terrestris, subspecies, biosystematics, enzyme electrophoresis, male
appendages, intermediates, clines.
W. Landman, Noorder Dierenpark/Zoo, Emmen, The Netherlands, L. Oudman, Department
of Genetics, University of Groningen, P.O. Box 14, 9750 AA Haren, The Netherlands (adress
for requests for reprints); M. Duijm, Department of Zoology, University of Groningen; present
adress: Onnerweg 41, 9751 VB Haren, The Netherlands.
INTRODUCTION
Ephippiger terrestris (Yersin, 1854) occurs in
the French Alps, in the neighbouring Italian Alps
and also around the Italian-Swiss lakes. Nadig
(1980) argued that E. bormansı (Brunner von Wat-
tenwyl, 1882) has to be regarded as a subspecies
of E. terrestris and described a new subspecies,
E. t. caprai. Furthermore, he found intermediates
between E. ¢. terrestris and E. t. bormansi in a
broad area in the department Alpes Maritimes
(Fig. 1). Duijm and Oudman (1983) confirmed
these results; they showed that between bormansı,
caprai and terrestris no mating barriers occur. Mat-
ing attempts between these terrestris subspecies
and E. ephippiger failed, however, apparently due
to differences in cerci and epiproct in the male
and the place of the copulatory sockets in the fe-
male partner. Hartley and Warne (1984) con-
firmed these findings by measurements of male
cercal reach and the corresponding female inter-
socket distances.
In this study morphometric characters are in-
183
vestigated, primarily to consider the relations be-
tween the three subspecies mentioned and the po-
sition of intermediates, mainly from a transect near
Tende. Allozyme analysis is used to get an in-
dependent character set. The allozyme frequencies
in E. terrestris are compared with those in E.
ephippiger vicheti, the latter being chosen as it
occurs sympatrically with E. ¢. bormansi at site
14 near the Lago di Como.
MATERIALS AND METHODS
Collection. - Specimens were collected during
field trips in 1980 and 1981. For locations see
table 1 and figs. 1 and 2. In 1981 special attention
was given to a transect near Tende (sites 1-11),
where intermediates might be found according to
Nadig (1980) and our own preliminary observa-
tions in 1980. The collecting area varied from one
to several acres, depending on the population den-
sity. The animals were killed, measured, photo-
graphed and frozen in solid carbondioxide (-79° C)
within one day of capture.
184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Table 1. Collection sites of Ephippiger species.
No. Location Country/dept. Altitude (m) Date
Ephippiger terrestris
N. Mézel/Majastres F 04 1200 A) 20-viii-80
B) 24/28-viii-81
2 Col de Maure F 04 1350 24-viii-81
Br Gréolières/Plan de Peyron F 06 1100 29-viii-81
4. Col de Braus F 06 1000 30-viii-81
5 Col de Castillon F 06 700 31-viii-81
6 Cle. Scravaion It Liguria 820 3-ix-81
7 Tende F 06 830 A) 29/31-viii-80
B) 1/3-xi-81
8. Vievola 1 F 06 915 2-ix-81
9} Vievola 2 F 06 1050 2-ix-81
10. Col de Tende 1 F 06 1200 2/5-ix-81
11A. Col de Tende 2 F 06 1400 2-ix-81
11B. Col de Tende F 06 1350/1420 30-viii-80
12. Col de Tende 3 F 06 1600 2-ix-81
13: Vallone del Arma It Piemonte 1600 31-viii-81
14A. Naggio (L. di Como) It Lombardia 800 5/6-ix-81
15. Mt. Grona (Lago di Como) It Lombardia 1400 6-ix-81
We Collobrières F 84 135 26-viii-80
18. Fayence F 84 250 28-viii-80
19. Aiguines (Verdon) F 84 850 23-viii-80
Ephippiger ephippiger vicheti
14B. Naggio (L. di Como) It Lombardia 750 5/6-ix-81
16. Miglieglia/Mt. Lema CH Ticino 720 7-ix-81
Electrophoresis. — Preparation of samples, elec-
trophoretic techniques and the preparation of the
horizontal polyacrylamide gells were carried out as
in Van Dijk & Van Delden (1981). The following
eighteen enzyme loci were analysed: Alcohol de-
hydrogenase (Adh), Tetrazolium oxidase (To), two
Phosphoglucomutases (Pgm-2 and -3), three Este-
rases (Est-1, -2 and -5), Aldehyde dehydrogenase
(Aldox-2), two Hexokinases (Hk-2 and -3),
Fumarate hydratase (Fum), Xanthine dehydroge-
nase (Xdh), Glucose oxidase (Gluo-3), Aldolase
(Ald), Malic enzyme (Me), Isocitrate dehydroge-
nase (Idh), a-Glycerophosphate dehydrogenase
(a-Gpdh) and Glycerine dehydrogenase (Glydh).
A preliminary electrophoretic survey was made
in 1980 (Landman 1981). In this study ten of the
above mentioned loci were examined for E. terres-
tris from six localities (Nos. 1, 7, 11B, 17, 18 and
19). For polymorphic loci Mendelian inheritance
was assumed on the basis of enzyme band patterns
and checked by testing genotype frequencies for
every location for Hardy-Weinberg equilibrium.
Between populations Nei's genetic distances
were calculated (Nei 1975). From the matrix of
genetic distances a dendrogram was constructed
according to the UPGMA method (Sneath and
Sokal 1973). Principal component analysis was car-
ried out with the allele frequencies (arcsin Vp
transformed). The principal component scores of
each population were calculated and the first and
second represented in a diagram.
Morphology. - Five characters of body and legs
were measured in the field: body weight (BW),
body length (LB), length of pronotum (LP), length
of hind femur (LF) and length of fore tibia (LT).
Weight was determined by means of a Pesola
pocket spring balance (0 — 5 g in 0.1 g). Length was
measured with sliding calipers. In a number of
samples one character of the pronotum and one of
the head was scored. Nadig (1980) distinguished a
shield-shaped field medially in the metazona of the
pronotum and named it campus medianus; he
showed that the lateral limitation of this field is
sharper in E. t. bormansi than in E. t. terrestris and
E. t. caprai. Moreover, this field is smooth in E. t.
bormansi and more or less granular in the other
subspecies. We scored the surface texture with: 0 —
smooth, 1 - granular, and 2 — rough, but we did not
find the latter character-state represented in our
samples. Furthermore we paid attention to the
“fastigium verticis” on the head, the profile of
which, according to Nadig (l.c.) varies from very
LANDMAN, OUDMAN & Dum: Ephippiger terrestris 185
steep to receding, a steep fastigium being charac-
teristic for E. t. bormansi. We scored: 0 — steep, 1
- rather steep, 2 - receding.
Colours of pronotum and abdomen were also
noted in the field.
Cerci and epiproct of freshly killed specimens
were photographed with a Medical Nikkor (2 : 1).
From the projected diapositives drawings with an
end magnification of 40x were made. Measure-
ments were carried out on these drawings. To cha-
racterise the form of the epiproct the following
dimensions were measured (fig. 3A): apical, cen-
tral and basal width (EA, EC and EB, respectively),
lateral and medial length (EL and EM).
Fig. 3B shows the way in which the cerci were
measured: total length (CL = CA + CB) and its
division by the place of the inner tooth in length
of apex (CA) and basal length (CB), basal width
(CE) and apical width (CF) and finally, a measure
(CD) for the form of the outer lateral side of the
cercus, CD taken negative when this side is convex
and positive when concave.
Titillators were removed from the body and
measured with a stereomicroscope (20x) using an
ocular micrometer. Figure 3C shows the way in
which they were measured. TL indicates the total
length of the apical part of the titillator, TT the
length of the toothed part of it and TN the number
of teeth. The width of the apical part is measured
as TA, so that the ratio TA/TT indicates its slen-
derness.
Samples of collected specimens from all locali-
ties, including tips of abdomens of the specimens
used for electrophoresis, will be deposited in the
Entomological collection of the Institute for Tax-
onomical Zoology (Zoological Museum) in Am-
sterdam.
RESULTS
Allozymes
Eighteen loci were surveyed of which six were
polymorphic: Pgm-2, Pgm-3, Est-1, Est-2, Est-5
and Adh, with 3, 5, 2,4,2 and 2 alleles, respectively.
LI
7 x \
/ 4, Genève RL Be
7 i
3
Sr
SARAI
=. \__ Lago
Maggiore (x
Y
Mon
aco
E. terrestris
intermediates
Fig. 1. Collection sites and distribution of Ephippiger species in S. France, S. Switzerland and NW. Italy.
186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
a
a
H NE 7/1
1600m12 © y
ER 07
DI 71
1400 m 11 6,1)
al,
1200 m 10
“ Vievola 1
o‘
915 m
x
Fig. 2. Collection sites along the Tende transect (locations
7 -12) in Southeast France.
The allozyme frequencies are shown in table 2. The
genetic distances are summarized in table 3 and the
dendrogram calculated from these distances is
given in fig. 4.
The aggregate of all E. terrestris populations
| 5 |
|
CE-CF=CG
Fig. 3. Method of measurement of (a) epiproct, (b) cerci,
(c) titillators. For abbreviations see text.
appears to be well separated from E. ephippiger
vicheti (distance 0.1713). For E. t. capra: the separ-
ate position is also clear. The distance between E.
t. bormansi and E. t. terrestris is less. The popula-
tions along the Tende transect together form a
separate group.
The results of the 1980 study — as far as E.
terrestris is concerned - are summarized in table 4,
in which the distances for six populations are
given. In these data the sites 17 and 18 represent
with certainty the nominate form, whereas this
may be doubted with regard to the site 5 in the
former series. The 1980 and 1981 distances are
combined in the semi-geographical representation
of fig. 5. Comparison of these data appears to be
justified, because the distances between the popu-
lations 1 and 7, determined both in 1980 and 1981,
were quite similar: 0.0742 and 0.0785 respectively.
Principal component analysis was carried out for
14 populations of E. terrestris and for 18 allozymes.
Fig. 6 shows the relation between the principal
components 1 and 2. PC 1 and PC 2 together
explain 55% of the total variance. The distribution
of the 14 populations in this diagram is similar to
their position in the dendrogram (fig. 4), though
the methods used are quite different.
LANDMAN, OUDMAN & DUM: Ephippiger terrestris 187
Morphology
The measurements of body and legs are sum-
marized in table 5. The means of these measure-
ments are generally linearly interrelated as appears
from table 6. By way of illustration the relation
between body weight and body length is shown in
fig. 7. The E. t. bormansi populations are on aver-
age the smallest ones, even smaller than our minor
samples. Specimens of the nominate form, repres-
ented by the localities 17 and 18 and perhaps also
no. 5 (Col du Castillon) are the biggest ones. E. t.
caprai occupies an intermediate position and the
samples along the Tende transect are intermediate
between capra: and bormanst.
Table 7 shows that the mean body measure-
ments are negatively correlated with the altitude of
the collection site; especially for LT and LF the
correlation is rather close. This is illustrated in fig.
8 for the relation between LT and altitude. The
distribution of the samples is rather similar to that
in fig. 7, only the mean length of the tibia in Naggio
(No. 14A) is too short for the altitude of this site.
To compare the form of the epiproct in the
various populations only mean ratios are used (ta-
ble 8) and not absolute measurements, because
these strongly depend on body dimensions. The
variation in the ratios EL/EM and EA/EC is re-
presented in the diagram of fig. 9. The three sub-
species caprai, terrestris and bormansi appear to be
well separated. In the nominate subspecies EL/EM
is larger than in the other two subspecies. This
means that the hind margin of the epiproct is bent
genetic distance
inside so that the hind corners are protruding. The
ratio EL/EM in bormansi is the same as in caprai,
and significantly smaller than in terrestris, indicat-
ing that the hind corners are not prolonged and
may even be rounded. There is a clear difference in
the ratio EA/EC between bormansi and caprat: the
higher value in bormansi points to the often later-
ally projecting hind corners, whereas the low value
in caprai indicates the rounded hind corners of the
epiproct in this subspecies. The forma mznor has
the same high EL/EM ratio as the nominate form,
whereas its EA/EC ratio is higher. So in the mean
the hind corners are in minor somewhat more
laterally protruding than in the nominate form; in
this respect minor resembles bormansı.
Our measurements of the cerci are summarized
in table 9. Cercal dimensions are correlated with
the dimensions of other parts of the body. The
mean total length of the cercus (CL) is moderately
correlated with mean length of the tibia (r =
0.486), the measurements CD and CE closely (r =
0.916 and 0.949, respectively). Therefore we char-
acterize the cercal form by means of the ratios. In
fig. 10 the relative length of the apex (CA) is
compared with the relative deviation of a straight
outer margin (CD). The nominate form has the
longest apical part and a strongly concave outer
margin. E.t. bormansi has a shorter apical part and
a slightly convex outer margin. Er. caprai has a
rather short apical part anda slightly concave outer
side. The forma minor is intermediate between 7.
terrestris and t. bormansi, the populations along
0.2 01 0
IEEE |
16 > È 4
a à E.eph.vicheti
6 E.terr.caprai a
ena
8
9
- Tende transect o
10
ah
12 4
5 4
4 }E.t.terrestris? o
3 J
2 i
7 |E.t.t.f minor a
15 :
AR HE.t.bormansi e
Fig. 4. Dendrogram of Ephippiger terrestris populations, based on genetic distance, calculated from table 3 (1981 data).
188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Table 2. Allozyme frequencies of Ephippiger populations (n.m. = not measured).
Pgm-2 Pgm-3
No. Locality n a b c a b c d e
E. terrestris
1B. Mézel/Majastres 28 38 ‚62 - - - O1" -
2, Col de Maure 13 23 UT = = = 6) DI -
3 Gréolières 20 .50 40 .10 = DS) 38 38 =
4 Col de Braus 20 43 40 .18 - 39 DI A =
DI Col de Castillon 18 50 50 - Dy il - .07 -
6. Cle. Scravaion 23 .28 65 O7 - „96 04 = -
7 Tende 20 „10 „90 - - .60 .40 = -
8. Vievola 1 20 - 1.00 - — .70 25 - .05
9: Vievola 2 16 — 1.00 = - .69 Di = =
10. Col de Tende 1 15 = 1.00 = - 11 79-05 .05
il; Col de Tende 2 7 - 1.00 = - - 1.00 - -
12. Col de Tende 3 8 - 1.00 = - - .88 - 13
14A. Naggio 18 .69 il - - - 100 - -
19) Mt. Groma 10 .60 .40 - - .10 80 .10 -
E. e. vicheti
14B. Naggio Pedy OO) Zora 1) ENS: È
16. Miglieglia/Mt. Lema ER OO LOD hk he 2
Table 3. Genetic distances between populations of Ephippiger species.
E. terrestris
No. Localities 1B 2 3 4 5) 6 7
IB. Mézel/Majastres - .0102 .0243 .0276 .0555 .0822 .0785
2, Col de Maure - 0253 „0299 „0604 „0829 „0540
5): Gréolières - 0126 0304 0555 0627
4. Col de Braus = .0128 .0637 .0611
Ds Col de Castillon - ‚0535 0779
6. Cle. Scravaion = „0823
Do Tende -
8. Vievola 1
9. Vievola 2
10. Col de Tende 1
11. Col de Tende 2
12, Col de Tende 3
14A. Naggio
15: Mt. Grona
14B. Naggio
16. Miglieglia
LANDMAN, OUDMAN & DUIJM: Ephippiger terrestris 189
(Table 2, continued).
Est-1 Est-2 Est-5 Adh
a b a b c d a b a b
040 60 13 53 33 - 1.00 = 1.00 -
038 62 42 23 35 - 1.00 - 1.00 =
069 31 03 35 42 20 94 06 1.00 —
055 44 22 58 18 02 69 31 1.00 =
054 46 08 67 29 .62 38 1.00 =
1.00 — 10 37 33 - 94 06 80 20
1.00 - 95 03 03 - 1.00 = 97 03
92 08 72. 03 .25 - 78 22 1.00 =
89 11 79 - 21 - 88 12 1.00 —
96 04 37 03 .60 = 37 63 1.00 =
88 12 33 = 67 = 42 58 1.00 =
1.00 = 50 = 50 = 67 33 1.00 =
83 17 = 24 .76 = 79 21 1.00 =
1.00 = - 25 75) - 1.00 = 1.00 -
.67 33 - = .08 92 n.m. n.m.
44 56 - = = 1.00 n.m. n.m.
(Table 3, continued)
E.e. vicheti
8 9 10 11 12 14A 15 14B 16 No.
0784 0749 0672 0587 .0532 0297 0344 1946 1960 1B.
0534 0489 0555 0506 .0407 0454 0470 1993 2000 2
0566 0575 0639 0675 .0571 0278 0198 1464 1566 3
0508 0536 0628 0701 .0648 0462 0460 1744 1798 4
0570 0632 0840 0971 -0941 0729 0698 1678 1734 5
0771 0804 0929 1034 .0832 0852 0659 1153 1352 6
0078 0044 0570 0680 .0380 .0943 0729 2257 2455 7
- 0010 0367 0504 0316 „0895 0719 2362 2549 8
= 0437 0549 0332 0911 0722 2354 2527 9
= 0026 0072 0496 0532 2625 2853 10
= 0072 0429 0505 2682 2885 11
= 0442 0411 .2469 2701 12
= 0067 1831 „2010 14A
= 1770 2002 15
= 0034 14B
190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Table 4. Genetic distances, calculated from allozyme frequencies determined by Landman (unpublished) for six samples
of Ephippiger terrestris collected in 1980.
No. Location 17 18
17. Collobrières = ‚0233
18. Fayence -
19. Aiguines
LA. Mézel/Majastres
Col de Tende
We Tende
the Tende transect are approximately intermediate
between ¢. caprat and t. bormansi. Strikingly apart
is the position of the populations 4 and 5 owing to
their relatively very short apices.
The basal part of the cercus tapers somewhat
towards the apex; this tapering is measured by CG,
the difference between CE and CF. Figure 11 shows
the relative measure of tapering. This is least in
bormansi and strongest at the sites 4 and 5. The
nominate and the mznor form occupy an interme-
diate position, as does capraz. The three sites at the
Col de Tende (10 — 12) are intermediate between
terrestris and bormanst.
According to Nadig (1980) the apical parts of
the titillators vary in slenderness. Our observations
ALLOZYMFREQUENCIES
C 21980 ———
1981
f. minor
ZX
}
\
\
\
\
\
\
EN
048
terrestris
19 1A 11B 7
.0634 .0482 .0789 .0709
.0278 .0477 .0553 .0912
- .0480 ‚0518 1090
- .0341 0742
- 1247
(Table 10) confirm this statement; we determined
the ratio TL/TA for caprai at 5.8, whereas the
values for the three bormansi sites were 11.8, 10.5
and 12.2 (mean 11.5). The bormanst titillators are
therefore evidently more slender than those of
caprai. Those of the forma minor were similar to
bormansi. Along the Tende transect we found for
the sites 7 - 12 the ratios 6.4, 6.9, 8.05, 10.7, 11.0
and 10.7 respectively. So there is a clear gradient
between caprai and bormansi, respecially along the
first part of the transect.
The relative scores for the texture of the campus
medianus and the form of the fastigium verticis are
shown in fig. 12. With regard to the campus me-
dianus (fig. 12 A) the character states as specified by
bormansi
Dar 15)
‘053
caprai
Fig. 5. Semigeographical representation (not on scale) of genetic distances between adjacent populations of Ephippiger
terrestris (1981 and 1980 data).
LANDMAN, OUDMAN & DUIJM: Ephippiger terrestris 191
Nadig (l.c.) are clearly confirmed: the surface is
smooth on sites 13 and 14 and therefore in bor-
mansi, whereas the granular state is apparently
characteristic for caprai (sites 6 - 9). The situation
at the three sampling sites along the Col de Tende
(10 - 12) is intermediate with the hormansi state
prevailing at sites 11 and 12, whereas at site 10 the
caprai state predominates, indicating a rather steep
transition with regard to this character over an
altitude difference of 200 m.
The steepness of the fastigium verticis shows
also a gradual transition (fig. 12B). A steep fasti-
gium appears to be characteristic for E.t. bormansi
and a less steep or even receding one for E. t. caprat;
a transitional situation is found along the Tende
transect.
It has to be stressed that all the above morpho-
logical results have been obtained by using the
means of rather variable series of measurements;
the statements do, therefore, not necessarily apply
to the form of the various parts in individuals
belonging to the populations studied.
Figure 13 gives an overall picture of the colour
of the abdomen. The various shades noted were
here combined into three categories only, viz.
green, brown and grey. The predominance of grey
(in fact yellowish grey) in the nominate form (sites
17 and 18) is remarkable. At other places shades of
green and brown were most common, whereas
grey was rather rare. There appears to exist a
gradient along the Tende transect with the percen-
tage of green individuals changing from 100% to
45%. Table 11 gives a more detailed picture of this
situation by also giving the shades of green ob-
served. One sees that in capraz (site 6) only plain-
green specimens were found, whereas at the sites
along the transect other shades of green occurred,
though in small numbers. The decrease of the rel-
ative numbers of plain-green specimens along the
transect is obvious. In the same time the occurrence
of three rows of dark spots on the abdomen (a
bormansi character) increases along this transect,
as appears from Table 11. Very interestingly also
at site 3 a sizeable percentage of this character was
observed (32%, n = 28).
DISCUSSION
As typical representatives of the nominate form,
E. t. terrestris, we consider our samples from
Fayence (18) and Collobrières (17) from the lower
parts of the department Var. Here one finds the
biggest specimens. The epiproct usually has
markedly protruding hind corners. The cerci have
a long apex, a concave outer margin and the basal
part tapers somewhat towards the apex. Unfortu-
Fig. 6. Diagram of scores for Principal Components 1 and
2, based on allozyme frequencies for 14 locations of
Ephippiger terrestris.
BW-_3.45+0.18LB
r=0.981
LB (mm)
Fig. 7. The relation between mean body weight (BW) and
mean length of body (LB) in Ephippiger terrestris.
nately these populations are missing in our 1981
collections. The 1980 data indicate a distance D =
0.048 to the minor population of site 1.
Nadig (1980) described E. t. capra: as a new
subspecies from the Ligurian Alps, close to E. t.
terrestris, but differing in form of epiproct and
titillators and in colour; the fresh grassgreen body-
colour is an important character in living capra:
specimens. For the capra: population studied we
can confirm his results: epiproct with rounded
corners, titillators with rather thick apical part.
Moreover, our results indicate differences in the
form of the cerci: the apical part is on average
rather short and the outer margin less concave than
192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Table 5. Mean body dimensions with standard deviations and ranges for 17 localities of Ephippiger terrestris.
BW - Body weight, LB - length of body, LP - length of pronotum, LT - length of protibia, LF - length of hind femur.
No. Locality n BW (g) LB (mm) LP (mm) LT (mm) F (mm)
1A. Mézel 21 1.27 s= O13 28.0 + 1.36 7.40 + 0.42 7.75 = 0.28 151 se 059
Li = 15 DS = 29 68 — 8.7 is =82 el = 165
2. C. d. Maure 28 1.01 + 0.13 25.8 + 1.43 6.92 + 0.24 7.32 + 0.28 13.8 + 0.47
07 —=12 Dj DI 64 73 68 — 7.8 13,2 = 15.1
3. Gréolières 23 1.27 + 0.18 26.8 + 1.53 7.26 + 0.41 751114059, 15.0 + 0.60
0.8 — 1.6 22 — 2) 64 — 8.0 69 — 8.0 13.4 — 16.0
4. C. de Braus 20 1.82 + 0.16 30.2 + 0.87 7.93 + 0.43 8.36 + 0.26 16.3 + 0.53
LS = 22 DI SII TR AD 78 — 8.8 ISO = All
5. C. de Castill. 20 2.56 + 0.42 33.2 + 1.64 8.79 + 0.50 9.18 + 0.40 17.9 + 0.76
15953 Sl =S 81 — 10.0 O2 — 97 16.8 — 19.5
6. Scravaion 20 1.34 + 0.12 DS ac 1123) 7.59 + 0.40 7.89 + 0.38 SEEN
I = ILS DO = 3X0) 66 — 8.3 69 — 83 159 159
7A. Tende 18 125 as O2 26.6 + 1.04 7.44 + 0.40 7.76 + 0.35 15.3 + 0.71
LO — 15 Dy —= Ts} 6.8 — 8.2 Ti = 83 14.6 — 16.2
8. Vievola 1 21 1.04 + 0.15 25.7 + 0.86 7.19 + 0.51 7.39 + 0.32 14.4 + 0.68
08 —14 DA — AR 64 —82 68 — 79 12155
9. Vievola 2 19 1.02 + 0.10 25.5 + 0.90 7.10 + 0.26 7.39 + 0.33 14.4 + 0.45
OUD — 2 Dj I COG — 116) 68 — 8.2 154 151
10. C.d. Tende 1 17 1.01 + 0.13 24.7. ze 1.50 7.26 + 0.43 7.25 = 0.32 14.0 + 0.62
Oy 2 20 — 26 6.6 —8.1 CSS 1A. = Weil
11. Cd. Tende 2 19 1.00 + 0.10 24.0 + 0.70 7.12 + 0.28 7.01 + 0.24 13.5 + 0.59
O MZ 2 25 67) 70 GT = 79 IES) S422
12. C.d. Tende 3 23 1.07 + 0.09 24.9 + 0.97 VY ze (07 7.13 + 0.28 EVE OST
OD NZ Dj DG Gl a 67 76 128 = 153
13. Vall. d. Arma 24 1.01 + 0.09 25.0 + 0.86 6.81 + 0.39 6.97 + 0.24 13.4 + 0.45
Of} = il DI, = Al 65) YU 6 =74 12.0 — 14.4
14A. Naggio 23 0.79 0.10 Dj as lena 6.40 + 0.12 6.85 + 0.18 13.0 = 0.44
Oy IM mM) =25 62 — 6.6 06 72 1224
15. Mt. Grona 17 0.89 + 0.11 DAS =e IK) 7.15 = 0.43 7.07 + 0.41 13.4 + 0.58
Oy = Ill Dj =D 64 — 8.0 NM) — Tell 12.4 — 14.2
17. Collobr. 17 2.42 + 0.26 DO ae 1759 8.84 + 0.53 9.72 + 0.46 18.8 + 0.75
Dl SHD DI, = 36 8.002205 SLO) = 105 18.0 — 20.4
18. Fayence 25 2.41 + 0.28 33.0 + 1.37 8.27 + 0.48 9.41 + 0.46 IAS) ae (047/77
Ta = Hall 3036 WI = 9D 8.8 — 10.6 16.6 — 19.8
Table 6. Interrelations between mean body dimensions of Ephippiger terrestris (17 localities).
regression correlation
body weight/body length LB =0.18 BW — 3.45 r= 0.98
pronotum length/body length LB =0.18 LP X 2.22 r= 0.93
tibia length/{»mur length LF =0.49 LT + 0.45 r= 0.99
pronotum ler gth/femur length LF = 0.34 LP + 2.29 r= 0.95
Table 7. Relations between mean body dimensions (table 5) and altitude (H in km) for Ephippiger terrestris (17
localities).
LANDMAN, OUDMAN & DUM: Ephippiger terrestris
regression correlation
body weight/altitude BW =— 104 H+ 24 ne (0.75)
body length/altitude UB == 9/4 iat 3320 r = — 0.74
pronotum length/altitude PRE 816 r=— 0.69
tibia length/altitude it = — OEI 96 = OU
femur length/altitude LF =—3.65 H+ 188 r = —.0.82
Table 8. Ratios between some measures of the epiproct (cf. Fig. 3A) in Ephippiger terrestris.
No. Locality
5
Mézel
C. d. Maure
Gréolières
C. d. Braus
C. d. Castill.
Scravaion
Tende
Vievola 1
Vievola 2
C. d. Tende 1
11. C. d. Tende 2
12, C. d. Tende 3
13. Vall. d. Arma
14A. Naggio
157 Mt. Grona
17% Collobr.
18. Fayence
SYHIDQUBEYSR TE
11.0
LT =-0.00179 Alt. 9.6
80 r=-083
EB/EA
0.82 + 0.08
0.83 + 0.09
0.77 + 0.05
1.08 + 0.09
ES) as OM
19252013
113,230]
1.14 + 0.17
1.10 + 0.14
0.97 + 0.13
0.96 + 0.10
0.99 + 0.13
0.93 + 0.13
0.82 + 0.08
0.86 + 0.05
0.97 + 0.10
0.99 + 0.10
EA/EC
1.26 + 0.12
1.25 + 0.09
1.34 + 0.13
0.98 + 0.07
0.96 + 0.07
0.86 + 0.07
0.92 + 0.08
0.94 + 0.10
0.98 + 0.09
1.10 + 0.13
1.15 + 0.09
JA2EE 007
1.19 + 0.13
1.25 + 0.06
1.22 + 0.05
1.13 + 0.09
1.09 + 0.08
altitude (m)
=
1000
Fig. 8. The relation between
mean length of tibia (LT) and
altitude (H) in Ephippiger ter-
restris.
EL/EM
1.29 + 0.20
1.10 + 0.10
1.06 + 0.13
1.02 + 0.15
0.94 + 0.10
0.82 + 0.10
0.95 + 0.09
0.88 + 0.06
0.88 + 0.06
0.84 + 0.12
0.84 + 0.10
0.83 + 0.09
0.74 + 0.11
0.78 + 0.05
0.79 + 0.07
1.21 + 0.09
1.11 + 0.09
194 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
EA,
< EA» VEC ae
forma minor
EC 1.3 t.bormansi 2
eh A2 &
PIE
1.2 013
ol
120 a’
LE ge m'è tterrestris
inter-
mediates
A LOF n A
08
o7
0.9}
ao
t.caprai
( ) EL
oF JEM
Fig. 9. The form of the epiproct in Ephippiger terrestris:
relation between the ratios EA/EC and EL/EM (see
text).
15
B) x
CAL o”
A!
3
20} 1 È
x ee A
DI A
[18] ew
So A nt)
es 7
£ FRE ===
2 isk ol © CACA+CB-16.3-0.03°D/CA+CB
© 0” r=-0.028
Vv
>
= le
O
O?
10H
Di
ol \
ze iS Li zijden
5 0 5 10
CONVEX+- STRAIGHT —» CONCAVE CD
outer margin
Fig. 10. Diagram of the relation between the relative
length of the cercal apex (CA) and the value CD (see
text).
in the nominate form. Furthermore, we confirm
Nadig's description of the granular campus medi-
anus and the receding fastigium verticis in caprat.
Our allozyme analysis showed an evidently distinct
position of capra: with a genetical distance D =
0.068 for E. t. terrestris and D = 0.076 for E. t.
bormansi. Such distances are rather low, but still in
the normal range for subspecies, for which Menken
& Ulenberg (1987) give a general range of 0.02-
0.22.
Our study of three populations of E. t. bormansı
confirmed Nadig’s (1980) observations. The anim-
als were generally very small and the abdomen is
often marked with three rows of dark spots and a
lateral yellow band. The epiproct has in the average
rather small, laterally protruding, hind corners.
The cerci have a short apex with a straight or even
slightly convex outer margin. The basal part of the
cerci tapers only slightly towards the apex; this
character corresponds with Nadig's not or hardly
thickened cercal base. The apical part of the titil-
lator is slender. The campus medianus is smooth
and the fastigium verticis steep. In our allozyme
analysis the distance to the E. £. terrestris popula-
tions is rather small: D = 0.0439, but this is prob-
ably an underestimation because we did not deter-
mine the distance to the populations no. 17 and/or
18 that were unfortunately not included in the 1981
series.
The forma minor is obviously characterized by
small body dimensions. We consider our samples 1
and 2 as typical. Table 5 and fig. 7 show that these
samples are in the mean much smaller than the
other E. t. terrestris populations, but somewhat
larger than E. t. bormansı.
As mean body dimensions are closely correlated
with altitude for the whole range of populations
studied (table 7, fig. 8) these dimensions cannot be
used as a distinctive character state for minor.
Grandcolas (1986) offered a similar biometrical
analysis and found the same relation with altitude
for ten populations ranging from 390 to 1264 m.
However, we found some small morphological dif-
ferences: usually the hind corners of the epiproct
are somewhat more prolonged (fig. 8) and there-
fore more similar to bormanst. Furthermore the
cercal apex is somewhat shorter than in the nom-
inate form. In our 1980 allozyme analysis the ge-
netical distance to terrestris amounts to 0.048,
rather high for closely related local forms (fig. 5),
whereas our 1981 analysis indicates a distance to
bormansi of 0.039. Though these values are not
entirely comparable, we suspect some bormansi
influence in our minor populations. Apparently
gene flow both in terrestris and in bormansi direc-
LANDMAN, OUDMAN & DuyM: Ephippiger terrestris 195
Table 9. Mean dimensions of the cerci of Ephippiger terrestris, cf. Fig. 3B (one unit = 1/40 th mm).
No. Locality n CA CB CD CL
1B. Mézel 21 Enk WD) 52.6 + 4.22 2.40 + 1.85 67.1 + 4.00
22 Col de Maure 29 MAR? 50.3 + 437 2.83 + 1.26 GLS == YA
DI Gréolières 23 1391322162 54.6 + 4.02 0.85 + 1.31 68.4 + 4.28
4. Col de Braus 20 Dl aa Deo! 65.2 + 3.04 3.75 + 2:42 70.8 + 3.53
DI C. d. Castillon 20 5.0 + 3.05 69.7 + 4.10 Joelle) 74.9 + 4.79
6. C. Scravaion 20 85311266 5 SEO 2.90 + 1.42 66.9 + 3.20
Vs Tende 29 7.0 + 3.41 56.1 + 2.34 222465 63.1 + 5.04
8. Vievola 1 19 10.6 + 2.93 DAN]? 1162 1.63 64.3 + 3.63
9, Vievola 2 19 1112162 55.2 + 4.34 0.24 + 1.29 66.1 + 4.59
10. C. d. Tende 1 17 IDEA 61.6 + 3.44 = 206,202] 74.1 + 3.86
ILA. Cd. Tende 2 20 IE OPES 21 60.4 + 3.23 MOLA 71.4 £ 3.53
122 C. d. Tende 3 23 1097223113 60.8 + 4.83 9122 TALS) ss)
15? Vall. del Arma 25 13-82-05 1] 66.2 + 6.22 — 2:90 1:01 74.0 + 4.50
14A. Naggio 23 11.6 + 1.69 55.6 = 2167) — 2.83 + 1.14 CHE 2:62
15 Mt. Grona 20 1922201 60.5 + 3.89 mea) HONTE SI
17% Collobr. al 182-2275 58.5 + 4.80 6.70 + 2.17 76.7 + 4.28
18. Fayence 25 18.2 + 4.20 OCSE 5.60 + 1.87 16,7 = 5.19,
CF CE CA / CL X 100 CD / CL X 100 CF / CE X 100 No.
32.8 + 1.80 ASS 7 Da 5 3.6 + 2.74 79.6 + 6.12 1B.
Mal ae.) 40.9 + 2.41 18.0 + 4.36 AS sate ZZ 80.2 + 4.50 De
329 se Di 41.0 + 2.84 202413192 lee WAX) 80.6 + 4.65 2°
DID ae 297) 45.2 + 2.49 8.0 + 3.46 531-2590 63.3 + 6.06 4.
30.8 + 2.48 49.2.2 227 6.7 + 3.92 7.7 + 2.68 62.6 + 4.50 DÌ
29.2 + 1.48 41.3 + 2.13 124270 4.4 + 2.16 70.9 + 4.31 6.
29.6 + 1.81 glassa, 10.9 + 451 1.8 + 2.58 71.9 + 4.62 Tks
29.4 + 2.41 41.2 + 1.63 15.5=25.41 1.8 + 2.47 71.5 + 4.43 8.
29.4 + 1.64 40.1 + 3.07 168-375 0.4 + 1.94 73.6 + 4.20 9.
Mey ae DAD 41.2 # 3.07 16.6 + 4.45 ZIO 78.9 + 5.63 10.
BV) ae I/O 40.1 + 2.76 15.3 + 4.16 — 2 as Dl 81.3 + 5.49 11A.
32.9 + 1.82 40.5 + 2.25 14.7 + 4.28 = Gas ey! 80.6 + 5.25 122
Wal as AO 39.1 + 3.60 18.8 + 4.23 AN SEO) 82.7 + 8.59 13:
ZINSEN GD, 34.3 1.79 SEE AA 91.5 + 6.40 14A.
DOED 38.8 + 2.43 WOT 22.2325) Sh) aN By 84.7 + 5.36 15
38.1 + 2.54 49.8 + 281 23.3 + 4.38 8.6 + 2.64 76.5 + 4.70 17.
35.0: 2.59 48.7 £ 3.82 23.8 + 5.10 Uvex Dy) 72.1 + 6.34 18.
tion is rather low, but nevertheless we think - with
Grandcolas (1986) - that the differences are too
small and too gradual for the designation mznor to
be used as an indication for any real taxonomical
unit. This term might only be used as a short-hand
for indicating the assembly of populations of ter-
restris living at higher altitudes, but without any
boundary.
The population near Gréolières (3) has many
small resemblances with the minor populations,
viz. form of epiproct, cerci and titillators. Its body
dimensions are in accordance with the altitude at
which it lives (fig. 8). In the allozyme analysis,
however, it is more closely related to populations
4 and 5 (figs. 4 and 5) than to the minor popula-
tions, to which the genetic distance is the same as
to bormanst. The occurrence of dark spot markings
in a significant number of individuals also indicates
a relation with bormansi. Nadig (1980) incorpo-
rated his specimens from Gréolières (Cheiron)
with the hybrids between terrestris and bormansı
on the basis of a great number of morphological
characters.
The populations 4 and 5 also offer some difficul-
ties with regard to their position. The form of their
epiproct is usually intermediate between caprai and
terrestris. Their cerci differ conspicuously from the
nominate form by the very short apex (fig. 10) and
the strongly tapering form of the basal part (fig.
11). In the allozyme analysis their distance to po-
196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Fig. 11. Relative measure of
tapering of the basal part of the
cercus.
50
pulation 3 is small (fig. 5), much smaller than to
the populations 6 and 7. Population no. 5 is judged
by Nadig (l.c.) as belonging to the nominate form.
We add the remark that these populations show
some differences with the more western popula-
tions from the terra typica and exhibit probably
some influence from caprai.
One of the aims of this study was to investigate
the populations along the Tende transect (sites 7-
12). Nadig (1980) collected specimens from the
Col de Tende which he judged to be hybrids be-
tween the subspecies terrestris and bormansi. Our
allozyme analysis shows all six populations along
the transect to be closely related (figs. 4 and 5). The
distances of this group to caprai and terrestris are
of the same order, those to bormansi are smaller.
In concordance with the geographical situation the
sites 7-9 form one subgroup and the sites at the
south side of the Col de Tende (10-12) another one,
the genetic distance of the latter group to bormansi
being significantly smaller. The form of the epi-
proct is intermediate between bormansi, terrestris
and caprai with the sites 7-9 closest to caprai and
the sites 10-12 closest to bormansi. In the form of
the cerci bormansi influences are generally evident,
but site 7 approaches caprai. With regard to the
slenderness of the apical part of the titillators a
gradient between caprai and bormansi was found
especially at the sites 7-9. The populations re-
sembled the bormansi character state. However,
this also appears to prevail in the minor popula-
tions. Nadig (Lc. fig. 36) showed the existence of
a continuous transition in slenderness from the
caprat- towards the bormansi-state with the terres-
CGE __6.59+0514CB
55
r=0.35
55 60 65 70
mean length of basal part of cercus (CB)
SEG t. see
oo CE
„en DS
A 77772
> En eee
a |] azz
I ee
pere > ui L
0 50 100% O RON SO TE 100%
E.t. bormansi
SI
A-CAMPUS MEDIANUS B-FASTIGIUM VERTICIS
Fig. 12. A (left): Distribution of the character states
granular and smooth of the campus medianus on the
pronotum in Ephippiger terrestris along the Tende tran-
sect. B(right): Distribution of the character states steep
- rather steep - receding of the fastigium verticis of the
head of E. terrestris.
LANDMAN, OUDMAN & Duy:
Table 10. The ratio TL/TA as a measure of the slender-
ness of titillators in Ephippiger terrestris.
No. Location n IEA.
DA Col de Maure 30 10.9 + 1.22
3. Gréolières 21 11.0 + 1.88
6. Scravaion 10 5.8 + 0.61
VÀ Tende 20 6.4 + 0.83
8. Vievola 1 20 6.9 + 1.03
9. Vievola 2 14 paco)
10. C. d. Tende 1 10 10974=t=M1826
11A. C.d. Tende 2 7 LOE 73
12. C. d. Tende 3 5 10.7 + 2.24
13% Vall. d. Arma 21 12.2 + 2.40
14A. Naggio 19 10924125
15 Mt. Grona 19 11.8 + 1.18
tris form at an intermediate and overlapping po-
sition. However, he did not relate these data to
geographical positions.
At sites 7-9 the campus medianus is granular, as
in caprai. At the sites along the Col de Tende this
state changes towards the smooth condition char-
acteristic for bormansı. A gradual transition is also
found with regard to the fastigium verticis: reced-
Ephippiger terrestris 197
ing in capraı and predominanty receding at sites 7-
9, with a gradual transition along the Col de Tende
sites towards the steep condition in bormanst.
With regard to the colour of the abdomen a
gradient was established in the occurrence of
(plain)-green and the abdominal marking with
rows of black spots.
Where Nadig (1980) doubted if capra was in-
volved in the origin of the intermediate popula-
tions and thought the mountain ridge forms a
sharp boundary for caprai, we think we have suf-
ficient evidence suggesting caprai influences exist-
ing in the region of intermediate populations, es-
pecially in the neighbourhood of Tende (where the
colour of the abdomen also resembles caprai), but
perhaps also in a wider area (viz. site 5).
The bormansi influence is quite clear at the Col
de Tende sites, but is — according to Nadig - much
more widespread. It is probable that this influence
is also present in the mzmor areas and is responsible
for the finding of supposed bormansi specimens as
mentioned in literature (viz. at the Col de la Cine).
Summarizing: it is fruitfull to consider Ephip-
piger terrestris - with Nadig - as a species occup-
ying an extensive area in SE France and N. Italy.
[ Jereen
= crey
[__]BRown
eOrange
eCarpentras
L
Fayence
De
CE
Ü
WA 144 15
\ Col de Larche ceva
( eCuneo
\
tà Col de Tende
©
/
3 4 5 ;
rc
& QC San Remo
D 3 nator:
eGrasse
Fig. 13. The colour of the abdomen in Ephippiger terrestris.
198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Table 11. Colour and abdomen marking in Ephippiger terrestris (percentages).
No. Locality n light plain
green green
6 Scravaion 32 = 100
7. Tende 25 5 93
8. Vievola 1 25 = 72
9 Vievola 2 24 4 92
10. C. d. Tende 1 28 = 64
11A. C. d. Tende 2 25 4 64
12, C. d. Tende 3 29 - 5)
13}, Vall. d. Arma 41 3 40
14A. Naggio 29 4 21
15. Mt. Grona 26 12 35
It is possible to distinguish the three subspecies
terrestris, caprai and bormansi, but only at the
outer margins of this area. All local populations
exhibit great variability in their characters, so that
it is often impossible to identify individuals to their
subspecies. No clear boundaries nor narrow hy-
bridisation zones exist between the subspecies, but
there is a large transitional field: in the centre of
the distribution area lies a quadrangular area (ap-
proximately 50 x 70 km) with intermediate popu-
lations, containing characters of all three subspe-
cies. Their influences manifest themselves often in
clinal characters, as was demonstrated by Nadig
and herein. No distinction between allopatric or
parapatric hybridization (e.g. Woodruff 1973) or
between primary and secondary intergradation ap-
pears possible, as long as no independent clues
about the history of the various taxa are available.
ACKNOWLEDGEMENTS
We thank the Uyttenboogaart-Eliasen Stichting,
Amsterdam and the Groninger Universiteitsfonds
for financial support. We thank prof. dr. W. van
Delden for providing laboratory facilities, for val-
uable discussion and continued interest. We are
also indebted to dr. H. van Dijk for instruction and
help with the electrophoresis techniques. We are
very grateful to dr. A. Nadig (Chur), who gener-
ously supplied us with information about his col-
lecting sites. Our thanks are due to H. J. de Vries,
who undertook the tedious work of measuring the
epiprocts. The assistance of M. Zijlstra and K.
Vrieling with computer matters is gratefully ac-
knowledged. We thank A. Decae, G. J. van Klinken,
H. Hiddingh, G. Overkamp, B. van Berkel, R.
Schoenmaker, J. van der Meulen, H. Steenbergen,
K. van Dijk, P. van der Hurk, H. Jansen, H. Von-
deling, B. Kruijt and L. Tromp for assistance in
collecting the insects. We are grateful to Dr. M. G.
dark greyish grey brown rows
green green of dark
spots
a 3 = x =
4 24 - - -
= = 4 2, =
4 15 4 15 11
8 - 24 29
- 14 = 55 49
- 12 8 25 —
- 48 7 11 52
- 39 16 - —
Ritchie for critically reading the manuscript and
for correcting the English. Furthermore we thank
Mrs. J. Poelstra for typing the manuscript and D.
Visser and D. van der Spek for making the draw-
ings.
REFERENCES
Dijk, H. van & W. van Delden, 1981. Genetic variability
in Plantago species in relation to their ecology. I.
Genetic analysis of the allozyme variation in P. major
subspecies. — Theoretical and Applied Genetics 60:
285-290.
Duijm, M. & L. Oudman, 1983. Interspecific mating in
Ephippiger (Orthoptera, Tettigonioidea). -
Tijdschrift voor Entomologie 126: 97-108.
Grandcolas, Ph., 1986. About the status of Ephippiger
terrestris forma minor. — Articulata 11: 299-305.
Hartley, J. C. & A. C. Warne, 1984. Taxonomy of the
Ephippiger ephippiger complex (ephippiger, cruciger
and centi) with special reference to the mechanics of
copulation. - Eos 60: 43-54.
Landman, W. L., 1981. Genetische afstand en verwant-
schap in de Ephippigeridae. - Internal report Depart-
ment of Genetics, University of Groningen. (unpub-
lished)
Menken, S. B. J. & S. A. Ulenberg, 1987. Biochemical
Characters in Agricultural Entomology. - Agricultural
Zoology Reviews 2: 305-360.
Nadig, A., 1980. Ephippiger terrestris (Yersin) und E.
bormansi (Br. v. W.) (Orthoptera): Unterarter einer
polytypischen Art. Beschreibung einer dritten Unte-
rart: E. terrestris caprai ssp. n. aus den Ligurischen
Alpen. - Revue Suisse de Zoologie 87: 473-512.
Nei, M., 1975. Molecular population genetics and Evolu-
tion. - North Holland Publ. Co., Amsterdam.
Sneath, P. H. A. & R. R. Sokal, 1973. Numerical Taxon-
omy. - Freeman, San Francisco.
Woodruff, D. S., 1973. Natural hybridization and hybrid
zones. — Systematic Zoology 22: 213-218.
Received: 4 January 1989
Revised version accepted: 29 May 1989
A TAXONOMIC REVIEW OF THE COLEOPHORIDAE
(LEPIDOPTERA) OF CHINA.
Contribution to the knowledge of the Coleophoridae, LIMI
by
GIORGIO BALDIZZONE
ABSTRACT
Baldizzone, G., 1989. A taxonomic review of the Coleophoridae (Lepidoptera) of China.
Contribution to the knowledge of the Coleophoridae, LIII. — Tijdschrift voor Entomologie
132: 199-240, figs. 1-120. [ISSN 0040-7496]. Published 1 December 1989.
The present taxonomic knowledge of the Coleophoridae of China is reviewed. Thirty-one
species are recorded here, of which fourteen species are described as new: Coleophora lucida,
C. buteella, C. caradjai, C. plurispinella, C. hoeneella, C. tibetana, C. cristata, C. alecturella,
C. sittella, C. falcipenella, C. yunnanica, C. denticulata, C. tuberculata and C. batangica. The
genitalia of C. seminalis Meyrick and C. summivola Meyrick are illustrated for the first time.
Two new synonymies are established: C. ëmmortalis Meyrick, 1922, a junior subjective synonym
of C. seminalis Meyrick, 1921 and C. nivifera Meyrick, 1930, a junior subjective synonym
of C. versurella Zeller, 1849.
Key words. — Coleophoridae; China; taxonomy; new species.
Dr. G. Baldizzone, Via Manzoni, 24, I-14100 Asti, Italy.
INTRODUCTION
The actual knowledge of the Chinese Coleopho-
ridae is very limited: up till now only the following
eight species have been reported from this vast
country: Coleophora neviusella Busck (Busck
1904), C. melanograpta Meyrick (Caradja & Meyr-
ick 1935), C. vibicigerella Zeller (= C. mandschu-
riae Toll), C. hsiaolingensis Toll (Toll 1942), C.
nivifera Meyrick, C. summivola Meyrick (Meyrick
1930), C. scioleuca Meyrick (Meyrick 1938) and
C. sinensis Yang (Yang 1983).
Judging from the important works by Falkovitsh
(1972b, 1974, 1975, 1976, 1977 and 1978), Fal-
kovitsh & Reznik (1980) and Reznik (1974, 1976
and 1977) the Coleophoridae are well represented
in Eastern Asia. They described many new species
from the rich material, representing about 100
species of Coleophoridae, collected by Z. Kaszab
in Mongolia. Their works on the fauna of Mongolia
and the region of Primorye (Southeast USSR)
form the basis of the study of Eastern Asiatic Co-
leophoridae. The fauna of Japan should also be
taken into consideration. Currently 31 species are
known from Japan (Baldizzone & Oku 1988a,
199
1988b) and many more new species, together with
a considerable number of species known from Eu-
rope, will be added to the Japanese list (Baldizzone
& Oku in press, in preparation). Finally, I have
just completed a paper on two new species from
Korea (Baldizzone 1989), whereas a list of the
twelve species known from that region will be
published later in collaboration with K. T. Park.
The aim of the present paper is to review the
taxonomic information on Chinese Coleophoridae
that is actually available. For that purpose I have
examined all material — wich is, unfortunately,
very little — that is kept in the museums of Europe
and the United States. The greater portion of this
material formed part of the well-known Höne col-
lection; the specimens that fortunately escaped
from the destructions of the second world war
have found their way to the Museum of Natural
History “Grigore Antipa” in Bucharest and the
Zoologische Forschungsinstitut und Museum
Alexander Koenig in Bonn. A small number of
specimens examined are kept in the Issiki col-
lection of the National Museum of Natural History
in Washington, whereas the majority of the Mey-
200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
rick types are in the British Museum (Natural His-
tory) in London. Unfortunately I have not been
able to study specimens recently taken in China or
material from collections within China. In the
material studied, 23 species were found in addition
to the eight mentioned above, making a total of 31.
Of these, 14 are here described as new.
I hope that this paper may form the basis for
future studies on the Coleophoridae of China, and
will enlarge our knowledge of this vast and com-
plicated family.
Concerning the systematics of the species
treated: I have followed Toll’s system, as ex-
pounded in his publications of 1952b and 1962.
Undoubtedly this system is due for a revision, based
on our present-day knowledge: according to my
opinion (and here I agree with Sattler & Tremewan
1978) the system proposed by Capuse (1971, 1975)
and Falkovitsh (1972a, 1987) is not useful and
complicates, instead of simplifies, the problems in
this family. Probably, instead of erecting a great
number of new genera, it is necessary to rearrange
the species within Coleophora into several species
groups. However, before one can succeed in such an
operation, the exact status of the species described
by the authors of the past should be assessed, and
modern methods should be employed to establish
the phylogenies of and the relations among the
different groups.
I have not prepared a key to the species treated
in this paper, as their number is too small in rela-
tion to what can be expected to be the ultimate
fauna of China. The 31 species treated here often
belong to groups that are, systematically speaking,
far from related, and it is to be expected that many
unknown species should be placed in between these
groups. In this case a comparison of the size of
Italy, where more than 240 species of Coleophori-
dae occur, with that of China is relevant.
Locality names are spelled according to the latest
edition (1986) of the “Times Atlas of the World,
comprehensive edition”. In this edition the Chi-
nese names are transcribed with the now widely
accepted pinyin romanization. Where the original
spelling on labels is deviating, this is added be-
tween square brackets. Information on the locali-
ties visited by Höne and cooperators is provided by
Niethammer (1963). A gazetteer of localities is
provided here as an appendix.
Abbreviations for museums:
BMNH British Museum (Natural History), Lon-
don, U.K.
Institute of Systematic and Experimental
Zoology PAS, Cracow, Poland
IZPC
INK Landessamlungen für Naturkunde, Karls-
ruhe, West Germany
MGAB Museum fiir Naturkunde der Humboldt-
Universitàt, Berlin, East Germany
RMNH Rijksmuseum van Natuurlijke Historie,
Leiden, Netherlands
USNM U.S. National Museum of Natural His-
tory, Smithsonian Institution, Washing-
ton, USA
Zoologisches Forschungsinstitut und Mu-
seum Alexander Koenig, Bonn, West Ger-
many
ZFMK
Checklist of the Coleophoridae of China
Coleophora Hübner, 1822
lucida n. sp.
neviusiella Busck, 1904
eurasiatica Baldizzone, in press
buteella n. sp.
melanograpta Meyrick, 1935
caradjai n. sp.
changaica Reznik, 1975
vibicigerella Zeller, 1839
mandschuriae Toll, 1942
hoeneella n. sp.
tibetana n. sp.
sinensis Yang, 1983
citrarga Meyrick, 1934
seminalis Meyrick, 1921
immortalis Meyrick, 1922 n. syn.
cristata n. sp.
scioleuca Meyrick, 1938
alecturella n. sp.
sittella n. sp.
versurella Zeller, 1839
nivifera Meyrick, 1930 n. syn.
hsiaolingensis Toll, 1942
vestianella (Linnaeus, 1758)
summivola Meyrick, 1930
bagorella Falkovitsh, 1977
yomogiella Oku, 1974
kurokoi Oku, 1974
falcipenella n. sp.
yunnanica n. Sp.
artemisiella Scott, 1861
denticulata n. sp.
tuberculata n. sp.
weymarni Toll, 1942
batangica n. sp.
ACKNOWLEDGEMENTS
On this occasion I wish to express my gratitude
to all those who have helped me, either by sending
material, or by giving me information. In particular
I would like to thank: Dr. Aurelian Popescu-Gorj
BALDIZZONE:
(MGAB), who with great diligence and patience
has traced all the Höne specimens in the storage
rooms of the Caradja collection, Prof. Dr. U. Roes-
ler (LNK), Dr. Klaus Sattler (BMNH), Dr. Don
Davis and Dr. Ronald Hodges (USNM).
A special word of thanks is due to my Dutch
friends Dr. Erik van Nieukerken (RMNH) and
Hugo van der Wolf (Nuenen) for their assistance
in realising this publication.
Coleophora lucida sp. n.
(figs. 1, 22, 40, 41)
Type material. — Holotype 8, China: Prov. Zhejiang
[Chekiang], West Tienmushan, 24.v.1932, H. Hone, ge-
nitalia slide Bldz 6892 (ZFMK).
Description. — Male (fig. 22) (Female un-
known). Wingspan 10 mm. Head (fig. 1), thorax
and abdomen nut-brown. Head with white scales
above the eye; labial palps white on inner surface
and suffused with brown scales on outer surface:
the second segment approximately half the length
of the third. Antenna with very thick tuft of nut-
brown short hairs at the base of the first segment;
flagellum ringed white and light brown. Forewing
with a uniform light nut-brown colour with a na-
creous lustre; a thin ochreous costal streak from
base to three quarts, fringes concolorous with
wing, but lighter. Hind wing dark pearl-grey;
fringes light yellow-grey.
Male genitalia (fig. 40). — Gnathos very big,
oval. Tegumen cylindrical, with two broad and
short arms. Transtilla narrow and stretched, trian-
gular. Valva short and thick, broader at the base
than at the top which is rounded. Valvula broad,
irregularly oval, covered with short bristles. Saccu-
lus with an almost straight ventral edge, the lateral
edge curved to the inside, forming an angle of
nearly 45° with the ventro-caudal angle. Aedeagus
slightly curved, long, dorsally more sclerotised. No
cornuti.
Structure of abdominal supports (fig. 41). — No
posterior lateral struts, transverse strut almost
straight, with a thicker distal edge. Tergal disks
(3rd tergite) oval, each with about twenty small
conical spines.
Diagnosis. — This species is difficult to place in
Toll's system, as it differs much from all known
Coleophora species. It may belong to the second
group because of its habitus and the form of its
tegumen and aedeagus. C. /ucida is closely related
to C. eteropennella Baldizzone & Oku, 1988a from
Japan. However, the following differences can be
observed: in C. /ucida the transtilla is narrower and
Coleophoridae of China 201
longer; the valva is narrower at the apex, the sac-
culus forms a more acute angle than that of
eteropennella, which is exactly 45° ; the aedeagus is
longer and more curved, and also the vesica is
longer; C. lucida is without any cornuti, whereas
eteropennella has a single big one, in the form of
a nail.
Biology. — Unknown.
Distribution. — Eastern China.
Coleophora neviusiella Busck, 1904
Coleophora neviusiella Busck, 1904: 45. Lectotype & (Se-
lected by Baldizzone, 1986: 140) China, [Prov. Shan-
dong], Yantai [Che-foo], 24.vi.1903, on apple, genita-
lia slide Bldz 7797 (USNM) [examined].
Coleophora eothina Falkovitsh 1974: 233 (Synonymized
by Baldizzone, 1986: 140).
Other material examined. — Paralectotype 9, same
data as lectotype, Mrs. Nevius, genitalia slide Bldz 7798
(USNM)
Diagnosis. — The species belongs to the 7th
group of Toll’s system. When publishing the syn-
onymy of C. eothina Falkovitsh, 1974 I dealt with
this species, and illustrated the female genitalia
(Baldizzone 1986).
Biology. — The species lives on apple (Malus
spp.) and Bramble (Rubus spp.). The case was
described by Falkovitsh (1974).
Distribution. — Eastern China and USSR, Pri-
morye.
Coleophora eurasiatica Baldizzone, 1989
Coleophora eurasiatica Baldizzone, 1989. Holotype, à,
China, Shanghai [Schangai, Prov. Kiangsu],
28.viii.1944, H. Hone (ZFMK) [examined].
Diagnosis. — The species, of which the female
is unknown, belongs to the 8th group of Toll’s
system and should be placed near C. algeriensis
Toll.
Biology. — Unknown.
Distribution. — China, Korea, Hungary.
Coleophora buteella sp. n
(figs. 3, 23, 44, 45)
Type material. — Holotype $, China: prov. Yunnan
[Provinz Nord-Yuennan], Lijiang [Li-kiang], 6.vii.1934,
H. Hone, genitalia slide Bldz 6887 (ZFMK). Paratype 1
8, data as holotype, but 30.vii.1935, genitalia slide Bldz
6884 (LNK).
Description. — Male (fig. 23) (Female un-
known). Wingspan 10-11 mm. Head (fig. 3) dor-
sally fulvous brown, laterally white. Thorax and
202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
abdomen light brown. Labial palps white on inner
surface, almost completely covered with brown
scales on outer surface; the second segment is about
twice the length of the third.
Antenna with a big tuft of short fulvous hairs at
the base of the first segment; flagellum ringed
white and dark brown. Fore wing uniformly ful-
vous brown, with a thin lighter costal streak;
fringes of the same colour.
Male genitalia (fig. 44). — Gnathos globular.
Tegumen cylindrical, narrowing at two-thirds,
with two short arms. Transtilla narrow, long,
rounded at the apex, where it is wider than at the
base. Valvula broad, the ventral edge rounded, co-
vered with short bristles. Sacculus heavily sclero-
tised, its ventral edge almost straight and its lateral
edge oblique; it is rounded towards the ventro-
caudal angle and ends in the dorso-caudal angle
with a crooked point which reaches the middle of
the valve. Aedeagus conical, long, simply formed.
Number of cornuti 4-5, in the form of needles of
different lengths, united at the base.
Structure of abdominal supports (fig. 45). —
Posterior lateral struts about half the length of the
anterior ones. Transverse strut straight and thin,
with a thicker distal edge. Tergal disks (3rd tergite)
about four times longer than wide.
Diagnosis. — This species belongs to the 9th
group of Toll’s system and should be placed in the
section of C. discordella Zeller. C. buteella can eas-
ily be distinguished from discordella by the shape
of the process in the dorso-caudal angle of the
sacculus, which is shorter and stouter, and by the
cornuti which are more numerous and shorter than
those of discordella.
Biology. — Unknown.
Distribution. — Only known from Yunnan, in
Southern China.
Coleophora melanograpta Meyrick, 1935
(fig. 24)
Coleophora melanograpta Meyrick, in Caradja & Meyr-
ick, 1935: 90. Holotype 6, China, Prov. Jiangsu
[Kiangsu], Longtan [Lungtan bei Nanking],
16.vi.1933, H. Höne, genitalia slide Bldz 8054
(MGAB) [examined].
Other Material examined. — 1 @, Prov. Zhejiang
[Chekiang], West Tienmushan, 28.viii.1932, H. Hone,
genitalia slide Bldz 6889 (ZFMK). 1 4, [USSR], Amur,
genitalia slide Toll 232 (MGAB).
Diagnosis. — The species belongs to the 16th
group of Toll’s system and should be placed
in the section of zelleriella Heinemann. This spe-
cies is treated by Baldizzone & Oku (in press).
Biology. — €. melanograpta lives on Oak
(Quercus sp.) and the larva occupies a pistol-
shaped case. Dr. Oku is studying the biology of the
species, following information I have found in the
Issiki collection in the USNM.
Distribution. — Eastern China, Korea and Ja-
pan.
Coleophora caradjai sp. n.
(figs. 11, 25, 48-54)
Type material. — Holotype @, China, Prov. Shandong
[Pr. Shantung], Taishan, mountain top [Gipfelhöhe], ca
1550 m., 29.viii.1934, H. Hone, genitalia slide Bldz 8053
(MGAB). Paratype 1 Q, data as holotype, but 12.viii.1934,
genitalia slide Bldz 6873 (ZFMK).
Description. — Imago (fig. 25). Wing span 14
mm. Head (fig. 11), thorax and abdomen white.
Labial palps white: the second segment about 2.5
times longer than the third. Antenna completely
white, with some long hairs at the base. Fore wing
with various shades of ochre towards the dorsal
margin and light brown towards the costal margin.
A thin white costal streak reaches the apex, where
the wing is completely brown. Fringes brown near
the apex and light yellow elsewhere. Hind wings
brown; fringes light brown-grey.
Male genitalia (fig. 48). — Gnathos small, oval.
Tegumen long, trapezoid, with two short and broad
arms. Transtilla very small and short, triangular,
rounded. Valva narrow and long, less curved at the
base than at the apex, club-shaped. Valvula very big
and sclerotised, triangular, covered with bristles.
Sacculus heavily sclerotised, curved, widening from
the base towards the dorso-caudal angle, where it
narrows into a small and sharp point which reaches
the middle of the valva. Aedeagus big, conical,
dorsally more sclerotised. Numerous cornuti of dif-
ferent lengths, united into a long, curved tress.
Structure of the abdominal supports (fig. 49). —
No posterior lateral struts. Transverse strut with a
straight proximal edge and a convex distal one.
Tergal disks (3rd tergite) about three times longer
than wide.
Female genitalia (fig. 53). — Papillae anales
small, oval. Apophyses posteriores almost twice as
long as apophyses anteriores. Lamella antevagina-
lis trapezoid, distal margin convex. It is deeply
excavated in the middle at the level of the ostium
bursae, which is wide, in the form of a cup. Infun-
dibulum long, funnel-shaped, with sclerotised me-
dian line as long as the ductus bursae. The ductus
bursae is very long, about 14 times longer than the
BALDIZZONE:
lamella antevaginalis: the upper part, which is 8
times longer than the lamella antevaginalis, is en-
tirely covered with many very small conical spines;
below that the ductus is transparent, with the ex-
ception of the median line, and ends in a wide
curve, at the end of which it is partially covered
with small spines towards the beginning of the
ductus seminalis. Bursa very wide, bag-shaped with
a leaf-shaped signum.
Diagnosis. — The species belongs to the 18th
group of Toll's system and should be placed in the
section of stramentella Zeller. C. caradjai n. sp. is
not difficult to identify because of its habitus — its
fore wings are two-colloured and because of the
following characteristics of the genitalia: In the
male, the gnathos is smaller and longer, the tegu-
men is wide and stout, the transtilla is longer, the
sacculus has a wider edge and ends in a shorter
point, there are fewer cornuti, united into a shorter
formation. With regard to the female genitalia it
can be observed that the lamella antevaginalis is
shorter, which is also the case with the apophyses
anteriores and posteriores. The ostium bursae is
wider and the spines of the ductus bursae are
smaller.
Biology. — Unknown.
Distribution. — The species has only been taken
in Eastern China.
Coleophora changaica Reznik, 1975
Coleophora changaica Reznik, 1975: 372.
Material examined. — 1 &, China, Prov. Shaanxi
[Sued-Shensi], Taibai Shan, Qin Ling [Tapaisan im Tsinl-
ing], 28.v.1935, H. Höne, genitalia slide Bldz 8512
(MGAB).
Biology. — The species lives on different species
of Artemisia.
Distribution. — China, Mongolia, Central Asia,
the Crimea, Jordan, Spain and Algeria.
Coleophora vibicigerella Zeller
Coleophora vibicigerella Zeller, 1839: 206.
Coleophora mandschuriae Toll, 1942: 291. Holotype 6,
China, Zalantun, in Nei Mongol Zizhiqu [Djalantun,
Manchuria, prov. Kirin], 1127’, 7.vii.1938 (IZPC)
[examined].
Biology. — The species lives on Achillea and
Artemisia sp.
Distribution. — China, Korea, North Africa and
Europe. |
Coleophoridae of China 203
Coleophora plurispinella sp. n.
(figs. 2, 26, 55-57)
Type material. — Holotype 8, China, Prov. Shandong
[Shantung], Taishan (1550 m.), 11.ix.1934, H. Hône,
genitalia slide Bldz 6888 (ZFMK).
Description. — Male (fig. 26) (Female un-
known). Wing span 13 mm. Head (fig. 2), thorax
and abdomen white. The second segment of the
labial palps is about twice the length of the third.
Antenna with a big bush of ochreous hairs at the
base of the first segment, which is white. Flagellum
light brown. Fore wing light ochreous, with a tri-
angular brown part along the radial veins. There
are silver lines along the costa, the medial, cubital
and anal veins. Fringes brown at the apex, yellow-
grey elsewhere. Hind wing light brown; fringes
light brown-grey.
Male genitalia (fig. 55). — Gnathos big, oval.
Subscaphium stout, narrowed at two thirds, with
two short and stout arms. Transtilla triangular,
pointed. Valva short, broad at the apex, narrowed
at the base, much curved. Valvula large, irregularly
trapezoid, the lateral edge sclerotised, covered with
bristles. Sacculus very much rounded, the edge
quite sclerotised, covered with bristles, on the lat-
eral edge below the dorso-caudal angle a big trian-
gular dent. Aedeagus short and conical, much
sclerotised basically and dorsally. A great number
of cornuti (more than 25), spines of different
lengths united into a long formation.
Structure of abdominal supports (fig. 56). — No
posterior lateral struts. Transverse strut thick, its
proximal edge straight and thin, the distal edge
thicker, convex. Tergal disks (3rd tergite) almost 4
times longer than wide.
Diagnosis. — The species belongs to the 18th
group of Toll's system and, because of the structure
of the male genitalia, should be placed in the sec-
tion of C. astragalella Zeller. The most obvious
differences are: in astragalella (fig. 58) the valva is
longer, the lateral process of the sacculus is shorter
and sharper, the number of cornuti is only 5-6,
whereas plurispinella has more than 25.
Biology. — Unknown.
Distribution. — The species has only been taken
in Eastern China.
Coleophora hoeneella sp. n.
(figs. 12, 27, 59-61)
Type material. — Holotype 9, China, Tibet, Batang,
Jinsha Jiang Valley [Im Tal des Yangtze] ca. 2800 m,
11.vi.1936, H. Höne, genitalia slide Bldz 8895 (MGAB).
204
Description. — Female (fig. 27) (Male un-
known). Wing span 15 mm. Head (fig. 12), thorax
and abdomen white. Labial palps entirely white:
the second segment is about three times longer
than the third. Antenna with a long, ochreous hair-
tuft at the base of the first segment; flagellum
ringed white and brown. Fore wing nut-brown, a
broad, white costal band, a long, white streak along
the cubital vein and a shorter one along the medial
vein. Fringes light grey-brown. Hind wing light
brown with light grey-brown fringes.
Female genitalia (fig. 59). — Papillae anales
broad and oval. Apophyses posteriores widenend
to a club-shape below the papilles, they are about
2.5 times longer than the apophyses anteriores.
Lamella antevaginalis trapezoid with many horiz-
ontal folds in the distal part. Lamella postvaginalis
trapezoid, very simple. Ostium bursae wide, oval.
Infundibulum cup-shaped. Ductus bursae massive,
oval, covered with very thick conical spines, form-
ing two bands along its first part, whereas the
second part is completely transparent, with the
exception of a small, curved part, which is weakly
spotted with chitine. Bursa wide, bag-shaped, with
a leaf-shaped signum.
Structure of abdominal supports (fig. 61). —
Posterior lateral struts about half the length of the
anterior ones. Transverse strut, almost straight,
with a proximal edge sclerotised in the middle and
a distal edge more sclerotised at the sides. Tergal
disks (3rd tergite) about 3 times longer than wide.
Diagnoss. — The species belongs to the 18th
group of Toll's system. Because of the structure of
the female genitalia, it should be placed in the
section of ditella Zeller. The following differences
with that species can be observed: the subgenital
plate of hoeneella is narrower and longer; the duc-
tus bursae is covered with spines over a shorter dis-
tance than that of ditella; the signum bursae is
smaller.
Biology. — Unknown.
Distribution. — Only known from the type lo-
cality.
Coleophora tibetana sp. n.
(figs. 13, 28, 62-65)
Type material. — Holotype g, China, Tibet, Batang,
Jinsha Jiang Valley [Im Tal des Yangtze] ca. 2800 m,
28.vii.1936, H. Hone, genitalia slide Bldz 8892 (MGAB).
Paratypes 2 @, data as holotype, but 1.vii.1936, genitalia
slide Bldz 8894, and 12.vii.1936, genitalia slide Bldz 8896.
Description. — Male (fig. 28) (Female un-
known). Wing span 18-19 mm. Head (fig. 13),
thorax and abdomen white. Labial palps white,
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
partially suffused with brown on outer surface: the
second segment is about twice the length of the
third. Antenna with a tuft of reddish hairs at the
base. Flagellum ringed white and light brown. Fore
wing ochreous in the dorsal and brown in the costal
half. Pearly-white streaks on the surface of the
wing: the longest along the costa from base to four-
fifths; two short, triangular streaks at the apex
along veins R3 and RS; a broad band along vein
M1, from the centre of the wing to the margin;
another broad streak along the cubital vein; the last
streak in the area of veins A(1+2). Fringes light
yellow-grey. Hind wing and fringes light yellow-grey.
Male genitalia (fig. 62). — Gnathos big, oval.
Tegumen narrow and long, cylindrical, with two
short arms. Transtilla small and short, triangular.
Valva narrow and long, narrower at the base than
at the apex, club-shaped. Valvula large, irregularly
trapezoid, covered with bristles. Sacculus small, its
ventral margin weakly curved and its lateral mar-
gin straight, in the dorso-caudal angle ending in a
sharp traingular point, which reaches the base of
the valva. Aedeagus short and conical, only dorsally
sclerotised. The cornuti are numerous, in the form
of spines of different lengths, united into a shape
that resembles a brush.
Structure of abdominal supports (fig. 63). —
Posterior lateral struts about half the length of the
anterior ones. Transverse strut very thick, its prox-
imal edge convex and its distal edge almost
straight. Tergal disks (3rd tergite) about 4 times
longer than wide.
Diagnosis. — The species belongs to the 18th
group of Toll's system, but it is difficult to place it
in any known section because of the very character-
istic genitalia. It may be close to C. canarupennella
Toll, a species of Iran, which can easily be distin-
guished from tibetana because of the following
characteristics: in canariipennella the aedeagus and
the valva are considerably shorter, the cornuti are
fewer in number, united into a long row, whereas
the lateral margin of sacculus is fully crenated.
Biology. — Unknown.
Distribution. — Only known from the type lo-
cality.
Coleophora sinensis Yang
Coleophora sinensis Yang, 1983: 107. Holotype 9, China,
Prov. Shanxi [Shansi], 13.vi.1979. (North-Eastern
College of Forestry, China). [not examined].
Diagnosis. — The species, described on the basis
of 21 specimens, belongs to the 25th group of Toll's
system and should be placed near C. laricella
Hübner.
BALDIZZONE:
Biology. — The species is recorded to live on
Larix principis-rupprechti Mayr.
Distribution. — Only reported from Shanxi and
Hebei provinces.
Coleophora citrarga Meyrick
(fig. 10)
Coleophora citrarga Meyrick, 1934: 460.
Material examined. — 1
27.vi.1946, leg. Issiki (USNM).
Q, Taiwan, Taihoku,
Diagnosis. — The species belongs to the 30th
group of Toll's system and should be placed in the
section of glaucicolella Wood. It is treated in a
paper on some species of Japan (Baldizzone & Oku,
in press). i
Biology. — Unknown.
Distribution. — Japan and Taiwan.
Coleophora seminalis Meyrick
(figs. 16, 29, 66-73)
Coleophora seminalis Meyrick, 1921: 189. Holotype 6,
Java, Pekalongan, van Deventer, Coll. Piepers-
Snellen, “M.525”, genitalia slide Bldz 9247 (RMNH)
[examined].
Coleophora immortalis Meyrick, 1922: 556. Holotype 4,
Fiji Islands, Lautoka, 24.iv. on flower of Amaranthus
paniculatus, W. Greenwood (BMNH) [examined].
Syn. n.
Material examined. — China, 2 6, Shanghai [Schangai,
Prov. Kiangsu], 19.viii.1942 and 21.viii.1943, H. Hone
(ZFMK).
Male genitalia (fig. 66). — Gnathos big, oval.
Tegumen trapezoid, considerably narrowed to-
wards three quarters, with two long and broad
arms. Transtilla broad and flattened, irregularly
oval. Valva large, very broad, its dorsal margin
curved. Valvula small, heavily sclerotised, irregu-
larly oval. Sacculus broad, characterised by two tri-
angular points at the angles: the point at the ven-
tro-caudal angle is longer than that at the dorso-
caudal angle. Aedeagus narrow and long, consisting
of two sclerotised bands, of which one is thinner
and sharp at the apex, the other thicker, with a
curved tooth at the apex in ventral position. Cor-
nuti numbering 6—7, of different lengths, united
into an irregular row.
Structure of abdominal supports (fig. 67). — No
posterior lateral struts. Transverse strut straight,
its proximal edge thicker than the distal one. Ter-
gal disks (3rd tergite) about twice longer than
broad.
Coleophoridae of China 205
Female genitalia (fig. 70). — Papilles anales
narrow and long. Apophyses posteriores about
twice the length of the anterior ones. Subgenital
plate trapezoid, its distal margin convex with some
bristles; it shows two folds, symmetrical with the
sides of the ostium bursae. Ostium bursae oval,
opening at three quarters of the subgenital plate.
Infundibulum tube-shaped, medially expanded.
Ductus bursae with a median line in its first half as
far as the central curve; ductus covered with conical
spines over a section about twice the length of the
subgenital plate. The remainder of the ductus is
almost transparent. Bursa oval, with a signum of a
singular shape: a small irregularly oval plate
crowned by an abrupt upper edge. This signum
varies considerably and can also have numerous
rounded spines.
Diagnosis. — The species belongs to the 30th
group of Toll’s system and might be placed in the
section of glaucicolella Wood. I have frequently had
the opportunity to observe considerable variations
in the colouring of the wings, and even more in the
genitalia: in the male, the valva and the valvula, the
triangular processes of the sacculus, the point of
the aedeagus and the cornuti; in the female the
form of the signum bursae.
Biology. — Meyrick’s descriptions were based on
specimens reared from different species of
Amaranthus.
Distribution. — Fiji Islands, Java, Eastern China,
Australia, New Guinea, Sumatra (material recently
examined by me.)
Coleophora cristata sp. n.
(figs. 5, 30, 74-77)
Type material. — Holotype 8, China, Prov. Zhejiang
[Chekiang], West Tianmu Shan [Tien-mu-shan],
8.ix.1932, H. Höne, genitalia slide Bldz 6888 (ZFMK).
Description. — Male (fig. 30) (Female un-
known). Wing span 13 mm. Head (fig. 5), thorax
and abdomen white. Labial palps white, suffused
with reddish scales on outer surface: the second
segment is almost half as long as the third. An-
tenna entirely white, with the exception of a short
tuft of reddish hairs at the base of the first segment.
Fore wing completely white, weakly streaked red-
dish along the veins, with some brown scales in the
apical area. Fringes white at apex and light brown
at the dorsal costa. Hind wing and fringes light
brown.
Male genitalia (fig. 74). — Gnathos globular.
Tegumen narrowed towards three quarters with
two long, flattened arms. Transtilla broad, irregu-
larly trapezoid, its distal margin curved and point-
206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
ed. Valva large, wider at the apex than at the base.
Valvula small, trapezoid, covered with bristles. Sac-
culus with curved ventral margin and straight
lateral margin: a small rounded tooth in the ven-
tro-caudal angle, and in the dorso-caudal angle four
triangular teeth of different sizes, having the shape
of a cock’s comb. Aedeagus long, forming two
sclerotised bands, the thin one ending in a sharp
apex, the thick one in a triangular point, sur-
mounted by a long, curved protuberance. There are
five cornuti, of which four are curved, in the form
of spines of different lengths, whereas the fifth is
quite long, together with the others forming a kind
of talon.
Structure of abdominal supports (fig. 57). — No
posterior lateral struts. Transverse strut with a
convex proximal edge and an almost straight distal
one. Tergal disks (3rd tergite) about twice longer
than broad.
Diagnosis. — The species belongs to the 30th
group of Toll’s system and should be placed in the
section of C. virgaureae Stainton. The two species
can easily be separated by the following character-
istics of the male genitalia: in C. cristata n. sp. the
transtilla is larger with a sharper point in the
dorsal margin; the sacculus is narrower and the
teeth in the dorso-caudal angle are shorter and
stouter: the aedeagus is longer and the dorsal pro-
tuberance is thicker and longer than that in
virgaureae; the cornuti are longer and have a dif-
ferent shape.
Biology. — Unknown.
Distribution. — Eastern China.
Coleophora scioleuca Meyrick
(figs. 4, 31, 78-81)
Coleophora scioleuca Meyrick, in Caradja & Meyrick,
1938: 20. Lectotype @ (here designated) China, | Prov.
North Yunnan], Lijiang [Likiang], H.6-34, genitalia
slide BMNH 24443 (BMNH) [examined].
Other material examined. — Paralectotype & (abdo-
men missing), data as lectotype but 10.vi.1934, H. Hone,
(MGAB).
Male genitalia (fig. 78). — Gnathos oval. Tegu-
men triangular, considerably narrowed towards
three quarters, with two long and flattened arms.
Transtilla straight, curved at the apex, which has a
point curved backwards. Valva long, slightly
curved, rounded at the apex, very oblique. Valvula
irregularly triangular, heavily sclerotised, covered
with long setae. Sacculus narrow, with a large,
triangular fold at the base; the lateral margin is
curved, heavily sclerotised, and the dorso-caudal
angle has two big, rounded teeth. Aedeagus long,
curved, forming two bands, more sclerotised dor-
sally, of which the thin one ends in a narrow, sharp
point, whereas the thicker one is broader at the
apex, which is curved and ends in a sharp point.
Only one cornutus, which is very long, curved, and
fine in the form of a needle.
Structure of abdominal supports (fig. 79). — No
posterior lateral struts; transverse strut thick, with
a convex proximal edge and an almost straight
distal edge. Tergal disks (3rd tergite) about five
times longer than broad.
Diagnosis. — The species belongs to the 30th
group of Toll’s system and should probably be
placed in the section of therinella Tengström. As
the female is not known, it is impossible for me at
the moment to give it a more precise position.
Biology. — Unknown.
Distribution. — Southern China, Nepal.
Coleophora alecturella sp. n.
(figs. 15, 32, 82-85)
Type material. — Holotype 6, China, Tibet, Batang,
Jinsha Jiang Valley [Im Tal des Yangtze] ca. 2800 m,
3.viii.1936, H. Hone, genitalia slide Bldz 8913 (MGAB).
Description. — Male (fig. 32) (Female un-
known). Wing span 11 mm. Head (fig. 15), thorax
and abdomen light brown. Labial palps entirely
white on inner surface and almost entirely suffused
with nut-brown scales on outer surface: the second
segment is about twice the length of the third.
Antenna without hairtuft at the base. Flagellum
ringed white and brown. Fore wing uniformly
pearly light brown. Fringes brown. Hind wing and
fringes uniformly pearly brown.
Male genitalia (fig. 82). — Gnathos oval. Tegu-
men trapezoid, with two long arms. Transtilla
large, flattened, oval. Valva short and stout, ob-
lique. Valvula small, triangular, heavily sclerotised.
Sacculus small, strongly curved and thick on the
ventral and lateral margin, which ends in the
dorso-caudal angle in a structure covered with
small, rounded teeth of different sizes. Aedeagus
stout, consisting of two sclerotised bands, of which
one is shorter with a triangular point, the other
longer, surmounted by a triangular tooth on the
dorsal half, and with a small, rounded tooth at the
apex. Cornuti 7-8, of different lengths, united into
a long formation, thicker at the base.
Structure of abdominal supports: (fig. 83). — No
posterior lateral struts. Transverse strut slightly
convex. thinner in the middle. Tergal disks (3rd
tergite) about twice longer than broad.
Diagnosis. — The species belongs to the 30th
group of Toll’s system and should, because of the
BALDIZZONE:
structure of the male genitalia, be placed in the
section of sternipennella Zetterstedt, near
monoceros Falkovitsh. C. alecturella can be distin-
guished by the following characteristics: the aedea-
gus has a triangular tooth, the transtilla is oval and
the cornuti are long.
Biology. — unknown.
Distribution. — Only known from the type lo-
cality.
Coleophora sittella sp. n.
(figs. 18, 33, 86-89)
Type material. — Holotype 8, China, Prov. Yunnan
[Nord-Yuennan], Lijiang [Li-kiang], 9.vii.1934, H. Hone,
genitalia slide Bldz 6900 (ZFMK). Paratypes 2 8, data as
holotype, but 19.viii.1934, genitalia slide Bldz 6893 and
26.vii.1935, genitalia slide Bldz 6875; 1 8, Shanghai
[Schangai, Prov. Kiangsu], 19.v.1932, H. Höne, genitalia
slide Bldz 6869 (ZFMK, LNK).
Description. — Male (fig. 33) (Female un-
known). Wing span 12-13 mm. Head (fig. 18),
thorax and abdomen nut-brown. Labial palps
white, suffused with light brown scales on outer
surface: the second segment about half the length
of the third. Antenna with some reddish hairs at
the base of the first segment; flagellum ringed
white and nut-brown. Fore wing nut-brown
streaked with white along the main veins; fringes
pearly light brown. Hind wing uniformly brown,
with pearly light brown fringes.
Male genitalia (fig. 86). — Gnathos oval. Tegu-
men big, triangular, considerably narrowed to-
wards three quarters, with two long and very broad
arms. Transtilla very large, irregularly trapezoid.
Valva short and stout, wider at the base than at the
apex, which is rounded. Valvula oval, narrow and
long, heavily sclerotised. Sacculus with curved ven-
tral margin and straight lateral margin, with many
small teeth of different sizes in the dorso-caudal
angle. Aedeagus with a rather complicated struc-
ture, consisting of two sclerotised bands of widely
differing shapes: the shorter one ends in a rounded
apex and is surmounted by a long, curved and
serrated horn in its dorsal middle, the longer one
is rather thick and ends in a curved and bifurcated
apex, with two rounded points. Cornuti numerous,
shaped like curved needles of different lengths,
united at the base into a long structure.
Structure of abdominal supports (fig. 87). — No
posterior lateral struts; transverse strut wide, al-
most straight, its proximal edge thicker in the
middle and the distal one thicker at the sides. Ter-
gal disks (3rd tergite) about three times longer
than broad.
Coleophoridae of China 207
Diagnosis. — The species belongs to the 30th
group of Toll’s system and should be placed in the
section of sternipennella Zetterstedt. Because of
the structure of the male genitalia it is probably
closely related to C. /unensis Falkovitsh and C.
pseudolinosyris Kasy. C. sittella n. sp. can easily be
distinguished by the following characteristics: the
valva is shorter; the transtilla is wider; the sacculus
is different, mainly because of the straight lateral
margin; the aedeagus is stouter with a differently
shaped tooth in the middle, and a bifurcate point
which does not occur in other species; the cornuti
are more numerous and longer.
Biology. — Unknown.
Distribution. — South-western and Eastern
China.
Coleophora versurella Zeller
Coleophora versurella Zeller, 1849: 352. Lectotype 9
(here designated) Poland, Glogów [Glogau], genitalia
slide BMNH 4417 (BMNH) [examined].
Coleophora nivifera Meyrick, 1930: 625. Holotype (&
according to original description, abdomen and hind-
wings missing when examined), China, Tibet Yadong
[Yatung], B.10000’, 6.viii (BMNH) [examined]
syn.n.
Material examined. — China, Shanghai [Schanghai,
Prov. Kiangsu], 4 3, 13 9, 2.vi, 30.vii and 10-28.viii.1936;
10.vii and 18-19.viii.1942, H. Höne (LNK, MGAB).
Biology. — The species lives on the seeds of
Atriplex sp., Amaranthus sp. and Chenopodium
sp.
Distribution. — Widely distributed over the en-
tire Palaearctic region and North and South Amer-
ica.
Coleophora hsiaolingensis Toll
(fig. 7)
Coleophora hsiaolingensis Toll, 1942: 296. — Holotype
8, China, Xiaoling [Manchuria, Hsiaoling, Prov.
Kirin], 13.viii.1939, genitalia slide Toll 837 (IZPC)
[examined]. Allotype 9, data as holotype, but
20.viii.1939, genitalia slide Toll 838 (IZPC)
[examined].
Biology. — Unknown.
Distribution. — The species is known from
China and Japan. Recently I have studied a male
from Siberia (MGAB).
Coleophora vestianella (Linnaeus)
Phalaena (Tinea) vestianella Linnaeus, 1758: 536.
208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Material examined. — 1 9, China, Prov. North Yun-
nan, Lijiang [Likiang] ca. 3000 m., 7.ix.1934, H. Hone,
genitalia slide Bldz 6871 (LNK), 2 3, China, Prov. Shanxi
[Shansi], Mian Shan [Mien-shan], upland plain [Obere
Höhe], ca. 2000 m., 6.viii.1937, H. Hone, genitalia slide
Bldz 8515, and 10.viii.1937 (MGAB).
Biology. — The species lives on the seeds of
Atriplex sp. and Chenopodium sp.
Distribution. — Europe, Asia Minor, Iran, Afgh-
anistan, China and Japan.
Coleophora summivola Meyrick
(figs. 14, 34, 90-96)
Coleophora summivola Meyrick, 1930: 625. — Holotype
6, China, Tibet, Gyangzé [Giantse] B.14500’ J. 28”,
genitalia slide BMNH 24442 (BMNH) [examined].
Material examined. — 3 9, China, Prov. North Yun-
nan, Lijiang, 22.vi., 4.viii.1934 and 17.viii.1935, H. Hone
(LNK); 1 9, Prov. North Yunnan, Dégén [A-tun-tse],
upland plain [obere Höhe] ca. 4500 m., 10.viii.1936, H.
Hone (MGAB); 3 4, 2 9, Tibet, Batang, Jinsha Jiang
Valley [Im Tal des Yangtze] ca. 2800 m, 6 vii, 16.vii, 4.viii,
14-15.viii.1936, H. Höne (MGAB).
Male genitalia (fig. 90). — Comparison with
parenthella shows that the lateral margin of the
sacculus has no big, rounded tooth, which in
parenthella is present on the inner side of the
dorso-caudal angle. The aedeagus of summivola is
formed by two sclerotised rods of almost equal
lengths, both with a triangular tooth at the apex,
whereas in parenthella one of the two rods is one
quarter shorter than the other, and only the longer
one has a tooth at the apex. The group of cornuti
in the form of a talon in summivola is longer than
that of parenthella. The male genitalia of C. ae-
quigesa Falkovitsh also resemble those of
summivola. I have had the opportunity to study the
holotype of aequigesa, a species from Mongolia,
kept in the Museum of Natural History, Budapest,
and I have observed the following differences,
which are obvious in the aedeagus: the aedeagus of
aequigesa is formed by two rods that are perfectly
symmetrical, smaller than those of summivola, and
they have no teeth at the apex. The male genitalia
of aequigesa have no ductus ejaculatorius, which
was probably lost during the preparation of the
genitalia.
Female genitalia (fig. 94). — Compared with
parenthella the subgenital plate of summivola is
rather longer and more chitinous; the ostium bur-
sae is smaller, opening closer to the distal edge of
the plate; the infundibulum is longer and wider
than that of parenthella and the section of the
ductus bursae which is covered with spines is also
longer.
Diagnosis. — The species belongs to the 30th
group of Toll’s system, and should be placed in the
section of vestianella (Linnaeus), near parenthella
Toll.
Biology. — Unknown.
Distribution. — The species has only been taken
in China.
Coleophora bagorella Falkovitsh
Coleophora bagorella Falkovitsh, 1977: 592.
Material examined. — 1 g, China, Prov. North Yun-
nan, Dégén [A-tun-tse], upland plain [obere Höhe] c.
4500 m., 13.viii.1936., H. Hone, genitalia slide Bldz 8517
(MGAB).
Biology. — Unknown.
Distribution. — China and Mongolia.
Coleophora yomogiella Oku
(fig. 8)
Coleophora yomogiella Oku, 1974: 254.
Material examined. — 2 4, China, Prov. North Yun-
nan, Lijiang [Likiang], 1.vii.1934, H. Höne, genitalia
slides Bldz 6886 and 6897 (LNK).
Biology. — The species lives on Artemisia prin-
ceps and Artemisia montana.
Distribution. — Japan, Korea and China.
Coleophora kurokoi Oku
(fig. 9)
Coleophora kurokoi Oku, 1974: 256.
Material examined. — 2 8, 1 Q, China, Prov. North
Yunnan, Lijiang [Likiang], 1/.vi, 7.1x.1934 and
25.vii.1935, H. Hone, genitalia slides Bldz 6898, 6870 and
6899 (LNK), 1 9, Prov. Zhejiang [Chekiang], West Tian-
mu Shan [Tien-mu-shan], 18.viii.1932, H. Höne, genita-
lia slide Bldz 6891 (LNK).
Biology. — The species lives on Chrysanthe-
mum morifolium var. sinense and Artemisia prin-
ceps.
Distribution. — Japan and China.
Coleophora falcipenella sp. n.
(figs. 6, 35, 97-100)
Type material. — Holotype 6, China, Prov. North
Yunnan [Nord-Yuennan], Lijiang [Li-kiang], 10.vii.1934,
H. Hone, genitalia slide Bldz 7080 (ZFMK).
BALDIZZONE:
Description. — Male (fig. 35) (Female un-
known). Wing span 15 mm. Head (fig. 6) and
thorax white, suffused with reddish-brown scales
on dorsal surface. Labial palps white, suffused with
reddish-brown scales on outer surface: the second
segment is about 2.5 times longer than the third.
Antenna without hairtuft at the base; flagellum
ringed white and brown. Abdomen light brown.
Fore wing reddish-brown with numerous white
streaks along the costa and the main veins. Fringes
pearly brown-yellow. Hind wing light brown;
fringes brown-yellow.
Male genitalia (fig. 97). — Gnathos oval. Tegu-
men narrowed towards three quarters, with two
long arms. Transtilla flattened and curved, club-
shaped. Valvula irregularly oval, covered with thin
bristles. Valva short and stout, narrower at the base
than at the apex. Sacculus with curved ventral mar-
gin, ending in the ventro-caudal angle, forming a
right angle; the lateral margin is almost straight
and ends in the dorso-caudal angle, which goes
beyond the dorsal margin of the valva; a big,
rounded tooth can be found on the inside of the
lateral margin of the sacculus. Aedeagus very long,
certainly one of the longest of the family of Coleo-
phoridae; it is formed by two sclerotised bands,
very asymmetrical, of which the longer has two
curves and ends in a scythe-shaped apex. The shor-
ter band is about half the length of the other. It has
a triangular tooth a short distance from the apex.
A single long and curved cornutus, needle-shaped.
Structure of abdominal supports (fig. 98). — No
posterior lateral struts. Transverse strut thick, its
proximal edge straight, the distal one convex. Ter-
gal disks (3rd tergite) about 5 times longer than
wide.
Diagnosis. — The species belongs to the 30th
group of Toll’s system, but because of the peculiar
shape of the male genitalia it is impossible for me
to place it with certainty into a section already
known. For the same reason this new species can-
not be mistaken for any other species already
known: the structure of the sacculus and of the
aedeagus allow an immediate identification.
Biology. — Unknown.
Distribution. — Only known from the type lo-
cality.
Coleophora yunnanica sp. n.
(figs. 20, 36, 101-108)
Type material. — Holotype ¢, China, Prov. North
Yunnan [Nord-Yuennan], Lijiang [Li-kiang], 9.vii.1934,
H. Höne, genitalia slide Bldz 7084 (ZFMK). Paratypes 3
Q, data as holotype, but 22.vi., 10.vii. and 17.vii.1934,
genitalia slides Bldz 6894, 7083 and 7081 (ZFMK, LNK).
Coleophoridae of China 209
Description. — Male and female (fig. 35). Wing
span 12-13 mm. Head (fig. 20), thorax and abdo-
men ochreous. Labial palps white, suffused red-
dish-brown on outer surface: the second segment
about twice the length of the third. The ochreous
antenna has no hairtuft at its base. Fore wing uni-
formly ochreous, streaked with reddish-brown
along the main veins and suffused with some scat-
tered dark brown scales, mainly towards the apex.
Fringes light pearly brown-grey. Hind wing light
brown-grey; fringes concolorous with those of fore
wing.
Male genitalia (fig. 101). — Gnathos oval. Teg-
umen considerably narrowed towards three quar-
ters, with two very long and broad arms. Transtilla
short and small, triangular. Valva short and stout,
rounded at the apex. Valvula long, narrow, heavily
sclerotised, covered with long bristles. Sacculus
broad, heavily sclerotised, with curved ventral mar-
gin, with a long, triangular protuberance in the
ventro-caudal angle and an irregular process in the
dorso-caudal angle, resembling a spiral tooth. Vin-
culum with a conspicuous formation in its middle,
resembling a conical tooth. Aedeagus long, formed
by two highly symmetrical sclerotised bands: the
shorter and thinner one narrowing from the base
to the apex, where it ends in a point; the longer one
widening from the base to the apex, where it forks
into a longer and a shorter protuberance. Only one
cornutus, big and long, broad at the base, talon-
shaped.
Structure of abdominal supports (fig. 102). —
No posterior lateral struts; transverse strut weakly
convex at its distal edge, which is thicker than the
proximal one. Tergal disks (3rd tergite) about
three times longer than wide.
Female genitalia (figs. 105, 106). — Papillae
anales small and oval. Apophyses posteriores about
twice the length of the anterior ones. Lamella an-
tevaginalis trapezoid, its proximal margin convex
and its distal one concave, with long bristles; it has
a small, triangular tooth in each of the two corners
of its distal margin, and two broad, symmetrical,
transverse folds in the middle. Ostium bursae oval,
narrow and long, opening at the distal margin, and
reaching halfway down the subgenital plate. Infun-
dibulum very long and sclerotised, in the form of
a narrow bag, about four times longer than the
subgenital plate. The ductus bursae is undoubtedly
the longest so far known in the family of the Co-
leophoridae: about twenty times longer than the
subgenital plate; in its first part it is covered with
tiny conical spines in two bands, over about twice
the length of the subgenital plate; the remainder of
the ductus bursae is transparent and shows many
210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
coils in its middle section. The bursa is of normal
size, bag-shaped, with a leaf-shaped signum.
Diagnosis. — The new species belongs to the
30th group of Toll’s system, but because of the
unique shape of the genitalia it is really impossible
for me to place it into a known section. The male
genitalia slightly resemble those of C. pandionella
Baldizzone (1988), a species from Siberia and those
of €. issikit Baldizonne & Oku (1988a), a species
from Japan but there are rather obvious differen-
ces, mainly in the aedeagus and the protuberances
of the sacculus. The genitalia of the female, how-
ever, resemble those of species in the section of
galbulipennella Zeller (= otitae Zeller), such as
microtitae Toll & Amsel and treskaensis Toll &
Amsel, but there structure of the subgenital plate,
the infundibulum and the enormous length of the
ductus bursae allow an immediate determination.
Biology. — Unknown.
Distribution. — Only known from the type lo-
cality.
Coleophora artemisiella Scott
Coleophora artemisiella Scott, 1861: 409.
Material examined. — 2 @, China, Tibet, Batang, Jin-
sha Jiang Valley [Im Tal des Yangtze] ca. 2800 m., 22.v.
and 13.vii.1936, H. Hone, genitalia slides Bldz 8903, 8904
(MGAB).
Biology. — The species lives on Artemisia sp.
Distribution. — So far the species was only
known from Europe. The specimens from the Jin-
sha Jiang Valley indicate a much wider Palaearctic
distribution.
Coleophora denticulata sp. n
(figs. 19, 37, 109-112)
Type material. — Holotype @, China, Tibet, Batang,
Jinsha Jiang Valley [Im Tal des Yangtze] ca. 2800 m.,
1.ix.1936, H. Hone, genitalia slide Bldz 8910 (MGAB).
Description. — Male (fig. 37) (Female un-
known). Wing span 14 mm. Head (fig. 19) brown
dorsally and white laterally. Labial palps white,
almost completely suffused with brown scales on
outer surface: the second segment is about twice
the length of the third. Antenna without hairtuft
at the base; flagellum ringed white and brown. Fore
wing uniformly pearly light brown, suffused with
scattered white scales towards the apex; a broad,
white band along the costa, with gradations to-
wards dorsum. Fringes chocolate-brown. Hind
wing uniformly brown-grey; fringes chocolate-
brown.
Male genitalia (fig. 109). — Gnathos small,
pear-shaped. Tegumen narrowed towards two
thirds, with two long arms. Transtilla short, irreg-
ularly triangular. Valvula oval, with thick ventral
margin, covered with long bristles. Valva small,
rounded at the apex. Sacculus broad, heavily sclero-
tised, its almost straight ventral margin extended
in the ventro-caudal angle into a triangular, ser-
rated formation; its lateral margin almost perpen-
dicular, serrated up to the dorso-caudal margin,
which ends in a big formation, curved and serrated
at the external and straight at the internal margin:
the point of that formation reaches beyond the
dorsal margin of the valva. Aedeagus long and
curved, consisting of two sclerotised bands, of
which one is narrower at the base than at the apex,
where it ends in a point, and the other is broader
at the base and ends at the apex in a big, triangular
tooth, serrated at the proximal margin. Numerous
cornuti of different lengths, united into a curved
formation resembling a tress.
Structure of abdominal supports (fig. 110). —
No posterior lateral struts; transverse strut convex,
thicker at the sides than in the middle. Tergal disks
(3rd tergite) oval, about twice as long as wide.
Diagnosis. — The species belongs to the 30th
group of Toll’s system and can be placed near C.
pseudociconiella Tolland C. hungariae (Gozmany),
because of the structure of its male genitalia. C.
denticulata can be distinguished by the following
characteristics: the rather small and short valva
and the serrated process of the dorso-caudal angle
of the sacculus, which is considerably bigger than
that of the other species.
Biology. — Unknown.
Distribution. — Only known from the type lo-
cality.
Coleophora tuberculata sp. n.
(figs. 17, 38, 113-116)
Type material. — Holotype @, China, Tibet, Batang,
Jinsha Jiang Valley [Im Tal des Yangtze] ca. 2800 m.,
3.viii. 1936, H. Hone, genitalia slide Bldz 8901 (MGAB).
Description. — Male (fig. 38) (Female un-
known). Wing span 14 mm. Head (fig. 17) white
laterally and reddish-brown dorsally. Labial palps
white, partially suffused with reddish-brown scales
on outer surface: the second segment is about twice
the length of the third. Antenna without hairtuft
at the base: the first segment is white, suffused with
brown scales dorsally; flagellum ringed white and
BALDIZZONE:
brown. Thorax and abdomen brown. Fore wing
light ochreous, with brown scales scattered along
the main veins. Fringes grey-brown. Hind wing
grey-brown. Fringes of hind wing brown.
Male genitalia (fig. 113). — Gnathos oval. Teg-
umen triangular, narrowed towards two thirds
with two long, flattened arms. Transtilla broad and
flattened, hatchet-shaped. In the middle of the
transtilla a sclerotised formation of an extremely
singular shape: a stalk widening into an inverted
cone, with lots of sclerotised spines of different
lengths in the widest part; this structure, whose
purpose is quite unknown to me, is highly remar-
kable and is absent in all other species of Coleopho-
ridae so far known. Valvula tiny and long, highly
sclerotised in an irregular form, covered with long
bristles. Valva long, its dorsal margin undulating,
the ventral one curved; wider at the base than at the
apex. Sacculus straight, heavily sclerotised, its ven-
tral margin curved, the lateral one straight: in the
ventro-caudal angle a small, triangular tooth; in the
dorso-caudal angle another bigger and longer
tooth, rounded at the apex, curved towards the
inside. Aedeagus consisting of two asymmetrical
sclerotised bands, wider at the apex than at the
base: they are slightly different in length, ending in
sharp points. Only one long cornutus with a wi-
dened base, in the form of a spine.
Structure of abdominal supports (fig. 114). —
No anterior lateral struts; transverse strut has a
thin proximal edge, slightly convex, and a distal
one, almost straight, laterally more sclerotised than
in the middle. Tergal disks (3rd tergite) about four
times longer than wide.
Diagnosis. — The species belongs to the 30th
group of Toll’s system and might be placed in the
section of C. adspersella Benander. C. tuberculata n.
sp. can easily be identified by the remarkable for-
mation in the middle of the transtilla, by the tooth
in the dorso-caudal angle of a sacculus which is
considerably shorter than in adspersella, by the
aedeagus consisting of bands without teeth at the
apex, and by a cornutus which is shorter than in
adspersella.
Biology. — Unknown.
Distribution. — Only known from the type lo-
cality.
Coleophora weymarni Toll
Coleophora weymarni Toll, 1942: 289. — Holotype 4,
China, Zalantun, in Nei Mongol Zizhiqu [Djalantun,
Manchuria, prov. Kirin] 1127, 10.v1.1938, genitalia
slide Toll 834 (IZPC) |examined].
Other material examined. — Paratype 9, data as ho-
Coleophoridae of China 201
lotype, but 26.vi.1939, genitalia slide Toll 835 (IZPC)
[examined].
Biology. — Unknown.
Distribution. — Only known from the type lo-
cality.
Coleophora batangica sp. n.
(figs. 21, 39, 117-120)
Type material. — Holotype g, China, Tibet, Batang,
Jinsha Jiang Valley [Im Tal des Yangtze] ca. 2800 m,
7.1x.1936, H. Höhe, genitalia slide Bldz 8909 (MGAB).
Description. — Male (fig. 39) (Female un-
known). Wing span 13 mm. Head (fig. 21), thorax
and abdomen entirely white. Labial palps ochreous,
with white scales mainly on inner surface: the sec-
ond segment about 1.5 times longer than the third.
The first segment of the antenna is white, suffused
with scattered brown scales, with a short, ochreous
hairtuft; flagellum ringed white and brown. Fore
wing reddish-brown, with broad costal streak, nar-
rowed towards two thirds and a white dorsal streak,
reaching the apex of the wing. The entire surface
of the wing has a pearly gloss. Fringes brown. Hind
wing brown-grey. Fringes brown.
Male genitalia (fig. 117). — Gnathos very small,
globular. Tegumen considerably narrowed at the
base of the socii, with two very broad triangular
arms. Transtilla united in the middle, broad and
triangular. Valvula tiny and long, heavily sclero-
tised. Valva very long and curved, at the base con-
siderably narrower than at the apex, on the dorsal
margin covered with bristles. Sacculus small,
heavily sclerotised, covered with long bristles, ex-
tending into a triangular form, obtuse at che lateral
margin, with two spines of unequal lengths in the
dorso-caudal angle. Aedeagus of medium length,
conical, consisting of two sclerotised symmetrical
rods, narrower at the base than at the apex, where
they are rounded. Numerous cornuti of different
lengths, united into a long formation resembling a
curved brush.
Structure of abdominal supports (fig. 118). —
No lateral posterior struts; transverse strut very
tiny, convex in its central part. Tergal disks (3rd
tergite) about twice as wide as long, covered with
short, very sharp, conical spines.
Diagnosis. — The species belongs to the 34th
group of Toll’s system, and because of the structure
of its male genitalia should be placed in the section
of C. unipunctella Zeller, a species from which C.
batangica can easily be separated by means of its
habitus, or by the following obvious differences in
the genitalia: the valva of batangica is narrower at
212 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
the base, longer and more curved; the transtilla is
bigger, with a large, central triangular widening,
which is not present in #nipunctella; the sacculus
of batangica is wider, its lateral margin longer, and
the spines in the dorso-caudal angle are close to
each other, whereas in wntpunctella they are far
apart; the aedeagus of batangica is shorter and
stouter, and the cornuti are longer and more
numerous.
Biology. — Unknown.
Distribution. — Only known from the type lo-
cality.
RÉSUMÉ
La connaissance taxonimique actuelle sur les Co-
leophoridae de la Chine est revue. Trente-et-un
espèces sont reconnues ici. Quatorze espèces sont
décrites comme nouvelles: Coleophora lucida, C.
buteella, C. caradjai, C. plurispinella, C. hoeneella,
C. tibetana, C. cristata, C. alecturella, C. sittella, C.
falcipenella, C. yunnanica, C. denticulata, C. tuber-
culata et C. batangica. Les genitalia des espèces
suivantes sont illustrés pour la première fois: C.
seminalis Meyrick et C. summivola Meyrick. Les
nouvelles synonymies suivantes sont établies: C.
immortalis Meyrick, 1922 est synonyme de C. se-
minalis Meyrick, 1921, et C. nivifera Meyrick, 1930
est synonyme de C. versurella Zeller, 1849.
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Received: 30 August 1988
Revised manuscript accepted: 12 February 1989
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di
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
APPENDIX
Gazetteer of place names
Names are listed alphabetically with their pinyin romanization. Province names are given in brackets after the
locality names. Sources for the spelling and coordinates were the “Times Atlas of the World, Comprehensive Edition,
7th ed., 1986” and the “Gazetteer of the People's Republic of China, 2 vols, 1979”.
Pinyin transcription Older transcriptions Coordinates
Batang (Tibet) !) Batang (also Pa-an) 30.02 N, 99.01 E
Dégén (Yunnan) A-tun-tse; Tehtsin 28.28 N, 98.48 E
Gyangzê (Tibet) Giantse, Chiang-tzu 28.57 N, 89.38. E
Jiangsu (prov.) Kiang-su, Chiang-su
Jilin (prov.) Kirin
Jinsha Jiang (river) Yangtze Kiang (part)
Lijiang (Yunnan) Li-kiang, Li-chiang 26.51 N, 100.16 E
Longtan (Jiangsu) Lungtan 32.10 N, 119.03 E
Mian Shan (mount) (Shanxi) Mienshan ca 37 N, 113 E
Nanjing (Jiangsu) Nan-ching, Nanking 32.03 N, 118.47 E
Nei Mongol Zizhiqu (prov.) Inner Mongolia auton. region
Qin Ling (range) (Shaanxi) Tsinling 34.00 N, 108.00 E
Shaanxi (prov.) Shensi
Shandong (prov.) Shantung
Shanghai (Shanghai) *) Schangai 31.13 N, 121.25 E
Shanxi (prov.) Shansi
Taibai Shan (mount) (Shaanxi) Tapaisan 33.57 N, 107.40 E
Taihoku (Taiwan) not traced
Taishan (mount) (Shandong) Tai-shan 36.20 N, 117.10 E
Tianmu Shan (mount) (Zhejiang) Tien-mu-shan 30.31 N, 119.36 E
Xiaoling (Jilin) 3) Hsiaoling not traced
Xizang Zizhiqu Tibet autonomous region
Yadong (Tibet) Yatung 27.29 N, 88.54 E
Yantai (Shandong) Yen-tai, Che-fou 37.30 N, 121.22 E
Yunnan (prov.) Yuen-nan, Yiin-nan
Zalantun (Nei Mongol Z.) Djalantun, Cha-lan-tun 48.00 N, 122.43 E
Zhejiang (prov.) Che-kiang, Che-chiang
Notes
!) Batang itself is situated in the province of Sichuan, but the nearby valley of the Jinsha Jiang forms the border with
Tibet. From the labelling with “Tibet, Batang, Im Tal des Yangtze” it follows that the specimens were probably taken
on the Tibet side of the river, or the border was not so distinct in those days.
2) Specimens from Shanghai are labelled as “Schangai, Prov. Kiangsu”. However, Shanghai forms nowadays a
province of its own.
5) There are various localities with the name Xiaoling (=Hsiaoling), but none was traced in the province of Jilin (=
Kirin). There is one located in the nearby province of Heilongjiang at 45.22 N, 127.17E.
BALDIZZONE: Coleophoridae of China 215
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Figs. 1-7. Heads of Coleophora. 1, C. lucida; 2, C. plurispinella; 3, C. buteella, 4, C. scioleuca; 5, C. cristata, 6, C.
falcipenella; 7, C. hsiaolingensis.
216 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
kurokoï: 10, C. citrarga; 11, C. caradjat; 12, C. boeneella; 13,
Figs. 8-13. Heads of Coleophora. 8, C. yomogiella; 9, G.
C. tibetana.
BALDIZZONE: Coleophoridae of China 23177
Figs. 14-21. Heads of Coleophora. 14, C. summivola; 15, C. alecturella; 16, C. seminalis; 17, C. tuberculata; 18, C. sittella;
19, C. denticulata; 20, C. yunnanica, 21. C. bantagica.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132
PPPPTIT TP CENA
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Coleophora SEP. 22, C. lucida, 8, holotype; 23, C. buteella, &, holotype;
holotype; C. plurispinella, 8, holotype; 27
DANG I 63 25, &
, C. boeneella, Q, holotype; 28, C. tibetana, 8, holotype; 29,
30, C. cristata, 6, holotype.
BALDIZZONE: Coleophoridae of China
2
9;
C. scioleuca, &; 32, C. alecturella, 8, holotype; 33, C. sittella, &, holotype; 34, C.
38, C.
Figs. 31-39. Coleophora spp. 31,
summivola, &; 35, C. falcipenella, 8, holotype; 36, C. yunnanica, 8, holotype; 37,
tuberculata, 6, holotype; 39, C. batangica, 8, holotype.
denticulata, &, holotype;
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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
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Figs. 40-42. C. lucida, holotype, slide Bldz 6892. 40, male genitalia; 41, abdomen; 42, detail of genitalia at high
magnification. Fig. 43. C. eteropennella, slide Bldz 8329, male genitalia, detail.
221
BALDIZZONE: Coleophoridae of China
14
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Figs. 44-47. C. buteella, holotype, slide Bldz 6885. 44, male genitalia, 45, abdomen; 46, detail of genitalia at high
magnification; 47, cornuti at high magnification.
222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 48-51. C. caradjai, holotype, slide Bldz 8053. 48, male genitalia; 49, abdomen; 50, detail of genitalia at high
magnification; 51, cornuti at high magnification.
<< er CC Sr
BALDIZZONE: Coleophoridae of China 223
Figs. 52-54. C. caradjai, paratype, slide Bldz 6873. 52-53, female genitalia, 54, subgenital plate at high magnification.
224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
99
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Figs. 55-57. C. plurispinella, holotype, slide Bldz 6872. 55, male genitalia; 56, abdomen; 57, detail of genitalia at high
magnification. Fig. 58, C. astragalella, slide Bldz 8709, male genitalia, detail.
BALDIZZONE:
Coleophoridae of China
225
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Figs. 59-61. C. hoeneella, holotype, slide Bldz 8895. 59, female genitalia; 60, subgenital plate at high magnification;
61, abdomen.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
[65]
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Figs. 62-65. C. tibetana; 62—63, 65, paratype, slide Bldz 8896; 64, holotype, slide Bldz 8892; 62, male genitalia; 63,
abdomen; 64, detail of genitalia at high magnification; 65, cornuti at high magnification.
BALDIZZONE: Coleophoridae of China 227
Figs. 66-69. C. seminalis, slide Bldz 6879. 66, male genitalia; 67, abdomen; 68, detail of genitalia at high magnification;
69, cornuti at high magnification.
228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 70-73. C. seminalis. 70—72, slide Bldz 7886. 70, female genitalia; 71, subgenital plate at high magnification; 72,
signum at high magnification; 73, slide Bldz 7888, signum at high magnification.
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Figs. 74-77. C. cristata, holotype, slide Bldz 6888. 74, male genitalia; 75, abdomen; 76, detail of genitalia at high
magnification; 77, cornuti at high magnification.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 78-81. C. scioleuca, lectotype, slide BMNH 24443. 78, Male genitalia; 79, abdomen; 80, detail of genitalia at high
magnification; 81, cornutus at high magnification.
BALDIZZONE: Coleophoridae of China 231
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Figs. 82-85. C. alecturella, holotype, slide Bldz 8913. 82, male genitalia; 83, abdomen; 84, detail of genitalia at high
magnification; 85, cornuti at high magnification.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
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Figs. 86-89. C. sittella, paratype, slide Bldz 6893. 86, male genitalia; 87, abdomen; 88, detail of genitalia at high
magnification; 89, cornuti at high magnification.
BALDIZZONE: Coleophoridae of China 235)
Figs. 90-93. C. summivola, slide Bldz 8897. 90, male genitalia; 91, abdomen; 92, detail of genitalia at high magnification,
93, cornuti at high magnification.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
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Figs. 94-96. C. summivola, slide Bldz 8911. 94, female genitalia; 95, subgenital plate at high magnification; 96, abdomen.
BALDIZZONE: Coleophoridae of China 235
Figs. 97-100. C. falcipenella, holotype, slide Bldz 7080. 97, male genitalia; 98, abdomen; 99, detail of genitalia at high
magnification; 100, cornutus at high magnification.
bo
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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 101-104. C. yunnanica, holotype, slide Bldz 7084. 101, male genitalia; 102, abdomen; 103, detail of genitalia at
high magnification; 104, cornutus at high magnification.
BALDIZZONE: Coleophoridae of China 237
Figs. 105-108. C. yunnanica, paratype, slide Bldz 6894. 105-106, female genitalia; 107, abdomen; 108, subgenital plate
at high magnification.
bo
Oo
(0 0)
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 109-112. C. denticulata. holotype, slide Bldz 8910. 109, male genitalia; 110, abdomen; 111, detail of genitalia at
high magnification; 112, cornuti at high magnification.
BALDIZZONE: Coleophoridae of China 239
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Figs. 113-116. C. tuberculata, holotype, slide Bldz 8901. 113, male genitalia; 114, abdomen, 115, detail of genitalia at
high magnification; 116, cornuti at high magnification.
240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
; ad te "x
4 .
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Figs. 117-120. C. batangica, holotype, slide Bldz 8909. 117, male genitalia; 118, abdomen; 119, detail of genitalia at
high magnification; 120, cornuti at high magnification.
CARABID FAUNA OF SOME TYPES OF FOREST IN THE
NETHERLANDS (COLEOPTERA: CARABIDAE)
THEODOOR HEIJERMAN & HANS TURIN
ABSTRACT
Heijerman, Th. & H. Turin, 1989. Carabid fauna of some types of forest in the Netherlands
(Coleoptera: Carabidae). — Tijdschrift voor Entomologie 132: 241-250, figs. 1-3, tables 1-3
[ISSN 0040-7496]. Published 1 December 1989.
Continuous pitfall trapping data were collected, consisting of year-samples of carabid beetles
from 48 sites in different types of forest located in several geographic parts of the Netherlands.
More than 18,000 carabids were caught. Numerical classification techniques were applied and
these revealed that clusters of sites, classified on their carabid species composition, could be
interpreted better in terms of their geographical position in the Netherlands, rather than in
terms of forest type. Moreover, the carabid faunas of the coniferous forests investigated, appeared
to be impoverished nor poor in species when compared with the faunas of deciduous forests.
Key words. - Carabidae; forest types; The Netherlands.
Th. Heijerman, Dep. of Entomology, Agricult. University, P.O. Box 8031, NL6700 EH
Wageningen, The Netherlands.
INTRODUCTION
If one were to try to record as many species of
forest carabids as possible from a single locality,
one would probably prefer to sample a mixed de-
ciduous forest, and certainly not a monotonous
coniferous plantation, where all trees are approx-
imately of the same age, size and are equidistant.
Den Boer (1963) studied the carabid faunas of
several types of forest in the province of Drenthe
(The Netherlands), and indeed concluded that the
carabid faunas of coniferous forests are only poor
sub-samples from those of dry deciduous forests.
Coniferous forests in Drenthe not only contain
lower densities of fewer carabid species, but also
lack the characteristic species.
Between 1835 en 1945, the Dutch forest area
increased from 170,000 to about 300,000 ha, which
is about 8.5% of the total area of the Netherlands.
This increase was due to the planting of various
coniferous tree species. The area covered with de-
ciduous forests thus decreased from 81% of the
total forest area to about 30%. At present about
80% of the total forest area is covered with conif-
erous trees, and only 20% with deciduous tree
species including scrubs. Deciduous forests often
alternate with coniferous plantations in the same
areas.
241
The patches of deciduous forests may therefore
be regarded as more or less isolated ecological “is-
lands”, surrounded by inaccessible areas. This hy-
pothesis would imply that not only agricultural
fields and meadows may function as barriers to the
exchange of carabid beetles between discontinuous
(spatially separated) habitats, but also that conifer
plantations may hamper dispersal to some extent.
Of course, this isolating effect would drastically
affect the distribution of stenotopic species, and of
species with poor powers of dispersal.
The possible effects of fragmentation and isola-
tion of the distribution of carabid species that differ
in their powers of dispersal, were recently studied
(Turin & Den Boer 1988).
In the present study the carabid faunas of several
forest types are described, the sampling sites are
classified, and the general applicability of the con-
clusion of Den Boer (1963), as mentioned above, is
tested. The results of our study seem relevant in
evaluating the possible role of conifer plantations
as ecological barriers, and therefore in interpreting
some of the results of the project of Turin & Den
Boer (1988).
Turin and Heijerman (1988) classified the
Dutch carabid species according to their association
with forest habitats, and where we speak of “true
242
forest carabids” we refer to the stenotopic and
eurytopic forest species as distinghuished in that
paper.
METHODS
Sampling technique and location of sampling
sites
We selected 14 sampling areas, distributed over
five geographical regions in the Netherlands (fig.
1). Each sampling area consisted of a forest area
containing patches of different types of woodland:
oak-birch, larch, spruce, and/or douglas-fir. Sets of
5 pitfall traps were placed in each of these different
forest types. In total 48 pitfall series were used,
covering 48 different sampling localities. Within
each pitfall series, traps were placed at a distance
of 6 to 10 m from each other. The traps were
provided with 4% formalin as a preservative. Ta-
ble 1 gives a listing of the sampling localities for
each sampling area.
The traps were emptied at irregular time inter-
vals during continuous operation for a whole year,
from spring 1983 to spring 1984. All those carabid
2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
specimens taken from the five pitfall traps of a set
together during a year, constitute a “year- sample”.
Statistical analysis
A data set was constructed containing the year-
samples of each species for each of the sampling
localities. Another data set was produced for only
those species that were classified as true forest
species by Turin and Heijerman (1988). In order to
simplify, visualize and to investigate the internal
structure of such data matrices and to obtain infor-
mation about the presence of clusters, two types of
multivariate analyses were performed on these
data sets. Both ordination and cluster analysis were
used. For information about the exact principles
and presuppositions of these methods, we will
refer to the literature.
Ordination and cluster analysis. - As an ordina-
tion technique we have used DECORANA to perform
Detrended Correspondence Analysis (DCA) (e.g.
Hill 1979a). This technique can be applied to inves-
tigate whether the sample sites can be arranged
= VIJLEN
= EPEN
= ES
= WIJLRE
= BEEK
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Fig. 1. Location of the forest
sampling sites in The Nether-
lands.
DO WOONDE © MNM
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VALKENHORST
HEIJERMAN & TURIN:
Carabid fauna of forest
243
Table 1. The 48 sampling sites, classified according to forest type and forest area. Given are the numbers of samples
(pitfall series) for the different kinds of forests and the serial numbers.
Geographical group forest area 1)
Southern part of 1. Vijlen TH+KA
Limburg 2. Epen TH+KA
3. Eys TH+KA
4. Wijlre TH+KA
Beek, Gelderland 5. Beek TH
Lievelde, Achterhoek 6. Lievelde EP
Southern Veluwe . Wageningen
7
8. Planken Wambuis KB
9. Warnsborn KA
10. Hoge Veluwe KA
11. Kemperberg KA
12. Renkum HT
13. Doorwerth HT
Brabant 14. Valkenhorst FS
forest type serial number
Total
oak-birch larch spruce douglas
2 2 3 - L1,2,5,7,8,11,13
i - 2 - L3,4,10
l I - - 16,9
- - - L12
1 2 - l B1,2,3,4
2 1 2 - A1,2,3,4,5
1 2 2 1 M18,19,20,21,22,23
5 2 2 - M1,2,3,4
- 2 1 - M5,6,9
- l 1 - M7,10
1 A te M8,11
- 2 2 M12,13,14,15
2 - - M16,17
l l l - NB1,2,3
12 16 16 4 48
1) Investigators: EP = E. Penterman; FS = F. Smulders, HT = H. Turin; KA = K. Alders, KB = K. Booij and
TH = Th. Heijerman
along one or more hypothetical axes. These axes
are constructed from the data matrix itself and may
or may not coincide with gradients of real environ-
mental variables. Subsequent consideration of the
ecological requirements of the species may be of
help to interpret the hypothetical gradient in terms
of real environmental variables. Two Way Indica-
tor Species Analysis was also carried out, using the
TWINSPAN program (Hill 1979b). This method can
be classified as a divisive, polythetic technique, and
groups sample sites into hierarchical clusters, on a
basis of the common presence of indicator species
and/or preferentials.
Diversity aspects. - The individual sample sites
were described in terms of (1) total number of
species and (2) total number of individuals. These
parameters were calculated for the total data ma-
tix and for the matrix containing the forest species
separately. They were used to investigate whether
or not the carabid species composition of the var-
ious types of coniferous forests constituted a poor
reflection of the communities of the deciduous
sites. The significance of differences found were
tested by the WILCOXON test.
RESULTS
The data matrices
The total carabid sample consisted of 18,522
specimens of 88 species (0.21 specimens trap*
day), twelve of them belonging to the stenotopic
forest species, ten eurytopic forest species (sensu
Turin & Heijerman 1988), and the others being
non-forest species. The total number of specimens
of true forest species was 17,281.
It is not practicable to present the complete data
set; table 2 gives the abundances of only the 22 true
forest species. The complete data set, however, is
available on request.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
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Fig. 2a. DCA-ordination (Detrended Correspondence Analysis). The sampling sites are marked according to their
geographical position in the Netherlands.
Statistical analysis
Ordination and cluster analysis. - The results of
the DCA-ordination are presented in fig. 2. The
analysis was performed using all species and all
sampling sites, and the original numbers of spec-
imens were transformed to their logarithms. In fig.
2a the sampling sites have been marked according
to their geographical position in the Netherlands.
In fig. 2b again the same sampling sites are shown,
but now the forest type is indicated. The positions
of sampling sites along the axes, representing hy-
pothetical gradients, obviously are not determined
by the type of forest habitat, but rather by their
geographical position. This seems particularly ob-
vious where it concerns the localities from Midden-
Nederland, Lievelde, Valkenswaard and the South-
ern part of Limburg (Zuid-Limburg), exclusive of
Wijlre. The Beek sample sites do not form a coher-
ent cluster, but are scattered among those from
Zuid-Limburg and Midden-Nederland. Wijlre con-
stitutes a very exceptional sampling site, being
situated in a douglas-fir stand very close to the river
Geul. Within Midden-Nederland clusters of forest
sites are rather obsolete. Clusters within the main
geographical regions are not indicated in the fig-
ures. Similar results were obtained using the forest
species alone, and are therefore not presented here.
Fig. 3a and fig. 3b show the results obtained by
TWINSPAN, based respectively on all carabid species
and on the forest species alone. Sampling sites are
marked according to their geographical position as
well as to the forest type. The main geographical
regions again appear to constitute more or less
concrete clusters; the Zuid-Limburg localities, ex-
clusive of Wijlre and inclusive of two Beek sites, are
grouped together in fig. 3a. Also the five sites of
Lievelde constitute a clear cluster. The Beek sites
are again distributed among the Zuid-Limburg and
the Midden-Nederland groups. The Valkenswaard
sites and the Midden-Nederland sites themselves
do not fall into clear clusters. Fig. 3b shows a rather
similar, though somewhat less clear picture.
From both the ordination and the TWINSPAN
analyses it can be concluded that the geographical
position, and not the forest type, is the important
factor determining the species composition of a
sampling site. It therefore seems impossible to
characterize the different types of forest in terms
of the presence of particular carabid species.
HEIJERMAN & TURIN: Carabid fauna of forest 247
] EN O (288)
FOREST TYPE: @ = Oak-BIRCH
A = LARCH
O = SPRUCE
150 -
= O = DoucLas FIR
À O
O Lo ©)
2° A
A
Sali 0% o ®
A ©) O ® 92
100 4 PR las ® + +
A A O A @ O
©)
O A
+
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504 N
L T An al T . EI 1
50 100 150 200 250
Fig. 2b. DCA-ordination, but the sites are now marked according to the forest type.
Diversity
The results of the diversity analysis are pres-
ented in tables 3a and 3b; in the first table all
species were considered and in the latter table only
the forest species were used in the calculations. In
these tables sites were grouped according to forest
type. The WILCOXON-test was used to test for dif-
ferences in the mean values of the diversity statis-
tics, between forest types. For all species together,
it can be concluded that there are more specimens
of carabids in deciduous forests as compared with
larch forests. Regarding the forest species, more
species of carabids were caught in deciduous forests
as compared with both larch and spruce forests.
Geographical clusters
Species abundance data can be used to define the
geographical clusters of sites that were detected.
We will not present all the indicator species and
preferentials as recognized by TWINSPAN for each
cluster, but only try to characterize the clusters
making use of the most important indicator species
as well as the abundance data from table 2.
Zuid-Limburg. From table 2 it can be seen that
Carabus coriaceus occurs on almost all Zuid-Lim-
burg sites, and also in all Beek sites. Thiele (1977)
summarized the data from a great number of inves-
tigations of carabids of the central European forest
communities of several associations of the Fageta-
lia and the Quercetalia (see his table 5). C. coriaceus
appears to be an important species of several forest
types in mountains. All Zuid-Limburg sites except
Wijlre, are situated at least 180 m above sea level,
and the sites near Beek are situated on a lateral
moraine (40 m above NAP [sea level]). However,
this species is also known from dikeslopes and
forests in the Betuwe (< 10 m above NAP). The
most dominant species in Zuid-Limburg are Abax
parallelepipedus, Pterostichus oblongopunctatus,
Carabus violaceus and C. problematicus, which are
also among the most abundant ones on many other
sites. C. problematicus is characterized by Thiele
(1977) as a species of mountains with no strict
affinity to a particular type of forest. The three
other species mentioned also occur in central Eu-
rope, independent of type of forest habitat. Rather
248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
striking is the absence of C. nemoralis, also a spe-
cies not characteristic of a specific forest habitat,
and common in all other geographical clusters.
Also N. rufipes, a species with a peak occurrence
in lowland Quercetalia, has not been caught. Geo-
graphical cluster 4 (fig. 3a) contains all Zuid-Lim-
burg sites but one (Wijlre) and two Beek sites, and
is defined by the indicator species C. coriaceus and
the non-forest species P. vernalis and P. madidus.
T. nitens and P. cristatus are restricted to Zuid-
Limburg. The last three species mentioned are im-
portant species of the Fagetum forests in central
Europe (Thiele 1977).
Midden-Nederland. Cluster 5 (fig. 3b) contain-
ing all but three Midden-Nederland sites, is defined
by TWINSPAN by the indicator species C. nemoralts.
This species is indeed less abundant in the other
geographical clusters except for the Beek sites.
Cluster 5 (fig. 3a) is characterized not only by C.
nemoralis but also by N. biguttatus and the steno-
topic forest species N. rufipes. True forest species
restricted to one or more Midden-Nederland sites
are A. brunnea and H. quadripunctatus. These two
OC
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bb
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Fig. 3a. Dendrogram showing the results of the
TWINSPAN cluster method, for all sampled species. G: sites
are marked according to their geographical position. F:
sites are marked according to the forest type. For mean-
ing of the symbols see fig. 2.
species are not menitoned by Thiele (1977) in his
table 5. N. biguttatus appears to be an important
forest species with no strict affinity to a particular
forest type.
Beek. The Beek sites do not constitute a homo-
genous geographical cluster; the sites intermingle
with Zuid-Limburg (through the occurrence of C.
coriaceus) and with Midden-Nederland (through
the occurrence of C. nemoralis). Also Beek has no
unique true forest species.
Valkenswaard. The Valkenswaard sites are
mainly characterized through the (almost) com-
plete absence of a number of forest species that are
abundant in the other areas: C. nemoralis, A. pa-
rallelepipedus and C. violaceus. There is one unique
forest species: L. spinibarbis, a species not menti-
oned by Thiele.
Lievelde. Again A. parellelepipedus is totally ab-
sent, as are C. violaceus and C. problematicus. Ca-
lathus rotundicollus is the most dominant species
and is much more abundant here than in the other
areas. C. rotundicollis is not mentioned by Thiele
as an important species of central European for-
i) Go Bed
I |
10 + |
|
: 009 | @400
SS ' AO Il
Ù I I
L__ 66 K 4
. —_ | 100888 | 440008
| 8000| 4400 |
10) 1 ! 1
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5 | 0000 I A © 1
TAO AA i
L 19 LA ‚A I
L__ eee | AOP
‚ee T NO
| ] E
‚Ae 1 Oe
ge 1 oe
1000 140
i i
az BE :@40
nee IN)
CL |
Fig. 3b. Dendrogram showing the results of the
TWINSPAN cluster method for the forest species alone.
Details as in fig. 2a.
HEIJERMAN & TURIN: Carabid fauna of forest 249
Table 3. Diversity statistics per forest type. 3a: calculated for all species, 3b: only true forest species. X: mean number,
SD: standard deviation. N-tot: total number of specimens per pitfall series, S-tot: number of species.
3a. N-tot S-tot H-tot
Forest type N x SD Xx SD x SD
oak-birch 12 DU? 491 15.6 5.4 1.5 0.3
larch 16 240 208 125 5.0 1.6 0.5
spruce 16 421 305 13.4 UL ile) 0.6
douglas 4 271 106 125) 4.7 1.6 0.1
3b. N-tot S-tot H-tot
Forest type N X SD X SD X SD
oak-birch 12 DO 495 IS) 1.8 1.4 0.2
larch 16 229 2a 7.8 1.8 1.4 0.3
spruce 16 394 368 74 22 1.0 0.4
douglas 4 268 105 9.8 Dl 1.6 0.1
ests, but this species is recorded as the most dom-
inant species in a mixed forest in Hall (DDR)
(Heckendorf et al. 1986) and in a mixed forest in
Foljuif (France) (Lecordier & Benest 1982). T. pla-
cidus is the only unique true forest species.
DISCUSSION AND CONCLUSIONS
Loreau (1984) studied the carabid communities
of a pine wood and a beech forest in Lembek (Bel-
gium). He found the beech forest to be inhabited
by more forest species compared with the pine
wood, to possess more characteristic species and to
have a higher species diversity. Inspection of the
values of the three components of diversity as cal-
culated by Loreau, however, reveals that differen-
ces are very small (l.c. Tableau V). Amongst other
things he finally concluded that the pine wood is
characterized by an impoverished carabid com-
munity. Recently Heckendorf et al. (1986) studied
the carabid communities of four types of forests
near Halle (DDR). The beech forest and the pine
forest communities were characterized as rather
poor in species as well as individuals.
These two examples confirm the findings of Den
Boer (1963) that coniferous forest types in general
can be regarded as marginal habitats for true forest
species, and that conifer plantations may indeed act
as barriers to the dispersal of carabids between
deciduous forest patches. These conclusions, how-
ever, are not really supported by our data. No
differences were found in the composition of the
carabid faunas of the forest types investigated, nor
are there any overall differences in the number of
species or the total number of specimens between
deciduous forests on the one hand and conifer for-
ests on the other. Our data strongly indicate that
the geographical position of the forest within the
Netherlands, constitutes a very important factor
determining the faunal composition. Also the idea
that coniferous forests may act as barriers that may
hamper dispersal of forest carabids between decid-
uous forests needs reconsideration.
ACKNOWLEDGEMENTS
We want to thank several colleagues who kindly
provided us with the data of their pitfall series (see
table 1): K. Alders, K. Booij, F. Smulders and the
late E. Penterman. Further we are grateful to P. J.
den Boer, P. H. van Tienderen and M. L. Luff for
their valuable suggestions. M. L. Luff also kindly
corrected the English.
250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
REFERENCES
Boer, P. J. den, 1963. Lebeort (Habitat)-Bindung einiger
Waldcarabidenarten in Drente (Holland) in Zusam-
menhang mit Waldtypus, Boden und Strukturelemen-
ten des Waldes (Paper read at Coenol. Symp., Zagreb
9/14-9 1963). — Publication (Comm. 115) of Biologi-
cal Station Wijster.
Boer, P. J. den, 1977. Dispersal power and survival. Ca-
rabids in a cultivated countryside (with a mathemat-
ical appendix by J. Reddingius). - Miscellaneous Pap-
ers Landbouwhogeschool Wageningen 14: 1-II,
1-190.
Heckendorf, Chr., A. Ruprecht, K. Schneider & F. Tietze,
1986. Zur Faunenstruktur (Coleoptera-Carabidae) in
Wald-Brachland Habitaten des NSG Lintbusch (158).
— Hercynia N. F 23: 72-82.
Hill, M. O., 1974. Correspondance analysis: a neglected
multivariate method. — Applied Statistics 23:
340-354.
Hill, M. O., 1979a. DECORANA: A Fortran program for
detrended correspondence analysis and reciprocal
averaging. — Ithaca New York.
Hill, M. O., 1979b. TWINSPAN: A Fortran program for
arranging multivariate data in an ordered two-way
table by classification of the individuals and attributes.
— Ithaca New York.
Lecordier, C. & G. Benest, 1982. Étude d'un écosysteme
forestier mixte, VI. Les Carabiques (Col.). - Revue
d'Ecologie et de Biologie du Sol 19 (1): 89-104.
Loreau, M., 1984a. Composition et structure de trois peu-
plements forestiers de Carabides. - Bulletin Académie
Royale de Belgique, Classe des sciences, Se Serie, 70:
125-160.
Loreau, M., 1984b. Les niches écologiques des Carabes en
milieu forestier, I. Composantes spatio-temporelles. —
Bulletin Académie Royale de Belgique, Classe des
sciences, Se Serie, 70: 333-379.
Thiele, H. U., 1977. Carabid Beetles in their environ-
ments. A study on habitat selection by adaptations in
Psysiology and Behaviour. - Zoophysiology and eco-
logy 10 (Berlin, Heidelberg, New York): I-XVII,
1-369.
Turin, H. & P. J. den Boer, 1988. Changes in the distri-
bution of carabid beetles in the Netherlands since
1880. II. Isolation of habitats and long-term time
trends in the occurrence of carabid beetles with differ-
ent powers of dispersal (Coleoptera, Carabidae). —
Biological Conservation 44: 179-200.
Turin, H. & Th. Heijerman, 1988. Ecological classifica-
tion of forest-dwelling Carabidae (Coleoptera) in the
Netherlands. — Tijdschrift voor Entomologie 131:
65-71.
Received: 19 August 1988
Revised version accepted: 1 August 1989
THE GENUS CRYPTUS FABRICIUS, 1804 OF THE CANARY
ISLANDS (HYMENOPTERA, ICHNEUMONIDAE)
G. VAN ROSSEM
ABSTRACT
Rossem, G. van, 1989. The genus Cryptus Fabricius, 1804 of the Canary Islands (Hymenoptera,
Ichneumonidae). — Tijdschrift voor Entomologie 132: 251-258, tables 1-7. [ISSN 0040-7496].
Published 1 December 1989.
Cryptus dianae Gravenhorst, 1829 solivagus subsp. nov. is described from Tenerife. A rede-
scription of the holotype of Cryptus obscuripes Zetterstedt, 1838 is given and the species is
acknowledged to be distinct. C. obscuripes impeditus subsp. nov. is described from Tenerife,
Palma and Gomera. The lectotype of Cryptus nigritarsis Kriechbaumer, 1894 from Tenerife is
redescribed. More specimens from Tenerife, only males, were found. Four new species are
described, viz., Cryptus praefortis from Tenerife and Gomera. Cryptus ultramondanus from
Lanzarote, Cryptus insularis from Fuerteventura and Cryptus furvus from Gran Canaria. Keys
to the Cryptus males and females from the Canary Islands are given.
Key words. — Cryptus, Canary Islands; new species.
G. van Rossem, Berkenlaan 25, 6711 RM Ede, The Netherlands.
INTRODUCTION
Two consignments of material from the Canary
Islands, received from Dr G. Ortega (Museo Insular
Ciencias Naturales, Tenerife) gave me the oppor-
tunity to study Canarian specimens of the genus
Cryptus Fabricius, 1804. Dr Kl. Horstmann (Zoo-
logisches Institut III, Würzburg) kindly offered me
to insert specimens from his collection and one
specimen from the Museo Insular at Tenerife. Dr
Horstmann also acted in my interest by asking for
the loan of the type material of Cryptus nigritarsis
Kriechbaumer at the Transvaal Museum at Preto-
ria (RSA). Dr Max Fischer (Naturhistorisches Mu-
seum Wien) sent the type specimen of Cryptus
verutus and Dr F. Koch (Museum für Naturkunde,
Berlin, DDR) put the type specimen of C. verutus
subsp. hierroensis at my disposal. Dr Roy Daniels-
son (Universitetets Zoologiska Institutionen,
Lund, Sverige) granted me the permission to study
the type of Cryptus obscuripes Zetterstedt and sev-
eral other specimens of that species and of Cryptus
dianae Gravenhorst as well.
With respect to the use of the name Cryptus the
author refers to Van Rossem (1987). Townes
(1970: 193) provides a description and a figure (fig.
165) of Itamoplex Forster, a junior synonym of
Cryptus Fabricius.
25]
In due time the collection Van Rossem will be
placed in the Rijksmuseum van Natuurlijke Histo-
rie at Leiden.
KEY TO THE CRYPTUS MALES OF THE
CANARY ISLANDS
The males of C. praefortis,
C. verutus are unknown.
C. furvus and
1. All femora orange coloured. Frons not concave.
Tyloids on flagellar segments 14-19. Inner or-
bits of face with broad ivory margin. Outer
orbit with narrow ivory margin. Face with
short pilosity. Mesoscutum widely and weakly
Puactured te TSE
ae Cryptus dianae solivagus subsp. nov.
Hind:femuniblatk ei Seen: 2
N |
. Face with conspicuous, long pilosity, the length
of which is reaching the width of mandibular
base int Re NN: 3
3. Front, middle femora and tibiae orange co-
loured. Hind femur fuscous, laterally ferrugi-
nous. Tyloids on flagellar segments
13(14)-17(18)(19). Face and gena with long
PreEyisetae sE rONSiNOMCONCIVCRE IA, EE
252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
N Cryptus nigritarsis Kriechbaumer
- Front and middle femora fuscous, brownish at
the extremity. Hind femur fuscous. Front and
middle tibiae brownish. Tyloids on flagellar
segments 12(13)-17(18). Face and gena with
long setae
EEE > Cryptus ultramondanus spec. nov.
. Basal area, first lateral area, areola and second
lateral area polished. Mesoscutum polished,
with fine punctation. Frons not concave. Ty-
loids on flagellar segments 15(16)-19
RE ER Cryptus insularis spec. nov.
— Basal area, first lateral area, areola and second
lateral area with sculpture. Punctation of me-
soscutum closer and coarse. Frons slightly con-
cave (not in a specimen from Palma). Tyloids
on flagellar segments 16(17)-19(20). Face and
gena with grey pilosity, but in length not reach-
ing the width of mandibular base
.... Cryptus obscuripes impeditus subsp. nov.
Ja
KEY TO THE CRYPTUS FEMALES OF THE
CANARY ISLANDS
The females of C. insularis and C. nigritarsis are
unknown.
IMÉlindemunoransecoloured non. 2
Hind femur black coloured
. Ovipositor exceptionally long, 0.96-1.0 of the
length of the front wing
DEE GERE AE Cryptus verutus Van Rossem
— Ovipositor of moderate length, 0.48-0.60 of
the length of the front wing (see remark on
page 253)
Beare Oe. SE Cryptus dianae solivagus subsp. nov.
. Ovipositor exceptionally long, 1.13 of the
N |
We
Table 1. Cryptus dianae dianae Gravenhorst
length of the front wing. Entire abdomen, with
allitergites) highlyspolishedi eis
Net a i di Cryptus furvus spec. nov.
= Ovpositorshorter ter 4
4. The apical transverse carina with strong apo-
physes. All abdominal tergites red in colour
and highly polished. Ovipositor 0.44-0.47 of
the lensth'ofithe front winless ee
NS. ORY OOD Cryptus praefortis spec. nov.
- The apical transverse carina without apo-
physes
5. Basitarsus of front leg 0.69 of the length of
front tibia. Ovipositor 0.86-0.98 of the length
of the front wing) AEREE RENO NOR ee
Passer NEN: Cryptus ultramondanus spec. nov.
— Basitarsus of front leg 0.58 of the length of
front tibia. Ovipositor 0.53 of the length of the
front wing (see remark on page 254)
.... Cryptus obscuripes impeditus subsp. nov.
Cryptus dianae dianae Gravenhorst
Cryptus dianae Gravenhorst, 1829: 545, Q.
Cryptus dianae dianae; Van Rossem 1969: 333-338. Com-
pare table 1.
Cryptus dianae dianae; Ortega & Baez 1980: 44-47.
Cryptus dianae solivagus subsp. nov.
Material. - Holotype: Tenerife, San Andres, 27.ii.1966,
10 m, leg. R. T. Simon Thomas, 1 8 (coll. Van Rossem).
Holotype label of the present author.
Description of male holotype. - Front wing 5.91
mm long. Palpi brown. Mandible black. Inner orbit
with ivory margin. Outer orbit with slight ivory
margin. Face black, pilosity short. Tyloids on flagel-
lar segments 14-19. Mesoscutum more widely
locality tyloids colour inner colour outer colour colour abdomen colour
orbits orbits femur 3 mandible
Netherlands 15-20 broadly ivory ivory marked orange postpetiole and terg. 2, 3 ivory; teeth
Grollo 18.v.1960 and 4 orange; rest fuscous fuscous
Denmark Löjt 15-20 broadly ivory ivory marked orange postpetiole and terg. 2,3, 4 ivory; teeth
Kirkeby vi.71 orange; rest fuscous fuscous
Italy Funes 15-21 broadly ivory ivory marked orange postpetiole and terg. 2, 3 ivory; teeth
20.vii.1968 and 4 orange; rest fuscous fuscous
Sverige Palsjò 15-19 broadly ivory ivory marked orange postpetiole and terg. 2, 3 ivory; teeth
Skane
and 4 orange; rest fuscous fuscous
VAN ROSSEM: Cryptus of Canary Islands 253
Table 2. Cryptus dianae solivagus subsp. nov.
locality tyloids colour inner colour outer colour colour abdomen colour
orbits orbits femur 3 mandible
Tenerife San 14-19 ivory margin narrow ivory orange tergites 2 and 3 orange; black
Andres 27.11.66 of face margin
rest fuscous
punctured than in continental specimens. All
femora orange coloured. First abdominal segment
black. Tergites two and three reddish, other ter-
gites black (third tergite orange in continental
specimens). Compare table 1 and 2.
Characteristics of the female. I did not see a
female from the Canary Islands of this subspecies.
Some important characters of the continental fe-
male follow here. Antenna with segments 4-7(8)
marked white, or without marking. Postanellus
slender 7.0-8.0 X as long as wide. Frons rather
concave. Inner and outer orbits with ivory margin.
All femora orange coloured. Abdominal segments
2-4 orange. Ovipositor in larger specimens
0.55-0.60 of the length of the front wing, 0.48-0.53
in small specimens.
Female. - Unknown.
Etymology. - "Solivagus” is the Latin for “stand-
ing solitary”.
Cryptus obscuripes Zetterstedt
Cryptus obscuripes Zetterstedt, 1838: 370.
Cryptus obscuripes, Horstman 1968: 311.
Cryptus dianae obscuripes; Van Rossem 1969:
336-337.
Cryptus dianae obscuripes; Ortega & Baez 1980:
47-49.
In my paper of 1969 I included Zetterstedt's
species, Cryptus obscuripes as a subspecies of
Cryptus dianae Gravenhorst. Reconsidering some
more or less stable features, I think this point of
view can not be maintained. It seems that
C. obscuripes represents a boreal - montane spe-
cies. It occurs in Scandinavia and Finland and also
it is known from mountainous localities in central
Europe.
Description of male holotype. - Labels: an orig-
inal label C. obscuripes 6, Talvig (Sverige, coll.
Zetterstedt). Holotype label of Horstmann (1966).
Front wing 7.33 mm long. Palpi, mandible, clypeus,
inner orbits, face below antennae, frontal orbit
next to antennal scrobes, upper inner orbit at OOL,
outer orbit (narrow), yellow to ivory in colour. Face
with grey pilosity. Frons, including scrobes, con-
cave. Upper frons with wrinkled sculpture. Anten-
nae, rather broken, with tyloids on flagellar seg-
ments 15-20 (left) and 15-19 (right). Prothorax
with rough sculpture and strong epomia. Mesoscu-
tum with rough punctures. Propodeum with coarse
wrinkled sculpture, both transverse carinae pres-
ent, the apical carina with weak apophyses. Meso-
pleurum with rough sculpture (greater part invis-
ible by pin). Front and middle coxae fuscous with
ivory spot. Hind coxae black, with conspicuous
setae. Front and middle femur brownish. Hind
femur brown with a ferruginous tendency. Front
and middle tibiae brownish with yellow streak,
slender. Hind tibia and basitarsus brown, the se-
cond, third and fourth tarsal segments white. Ab-
domen with petiole, seventh tergite and claspers
fuscous. Other parts orange in colour. Compare
table 3.
In all I have studied one other male from The
Netherlands, Ede, 28.vii.1970, leg. C. J. Zwakhals
(coll. Van Rossem). It closely agrees with the hol-
otype. Tyloids on flagellar segments 15-20 (on
20th segment very weak).
Description of female. - Sweden: Dir (Dalarna),
4.vii.1937, leg. A. Roman. Finland: 9, Fennia, Ks,
Salla, leg. Krogerus, 1936 (three specimens Uni-
versitetes Zoologiska Institutionen, Lund).
Front wing 6.66 mm long. Palpi fuscous. Entire
head black, except for ivory margins of upper inner
orbit, outer orbit and spot at OOL. Frons concave,
including antennal scrobes. There is some sculp-
ture around the median ocellus. OOL with micro-
sculpture. Antenna long and slender. Prothorax
with rough, longitudinal and wrinkled sculpture,
epomia strong. Mesoscutum polished, with scat-
tered, weak punctures. Propodeum with coarse,
wrinkled sculpture, both transverse carinae pres-
ent. Mesopleurum coarsely sculptured. All legs
brown. Abdomen with petiole fuscous, postpetiole
and all tergites orange. Ovipositor 0.53 of the
length of the front wing.
It seems to be unlikely that the species occurs in
the Canary Islands, though the mountainous char-
acter of that locality would make it acceptable.
I encountered three male specimens which I con-
sider to represent the species, although there are
some differences with the type, namely the posi-
254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Table 3. Cryptus obscuripes Zetterstedt
locality tyloids colour inner colour outer
orbits orbits
holotype Sverige 1 15-20 broadly ivory ivory marked
Talvig r 15-19
Netherlands Ede 15-20 broadly ivory ivory marked
28.vii.70
tion of the tyloids: 16(17)-19(20) versus
15-19(20) in the type; clypeus and face fuscous and
with ivory marking in the type; hind tarsal seg-
ments 2, 3 and 4 fuscous and ivory coloured in the
type.
Eventually the best thing to do is to propose a
subspecies for the Canary Islands specimens.
Cryptus obscuripes impeditus subsp. nov.
Material. - Holotype male: Tenerife, Montana Ber-
meja, l.iii.1966, 1500 m, leg. R. T. Simon Thomas (Coll.
Van Rossem). Holotype label of the present author. Par-
atypes: Palma, La Rosa (Mazo), 4.viii.1981, 1 4, leg. G.
Ortega (Mus. Ins. Cienc. Nat. HY 1347); Isl. Can. Go-
mera, Laguna Grande, 16.viii.1977, 1 8, leg. M. Baéz.
(Mus. Ins. Cien. Nat. HY 1339).
Description of male holotype. - Front wing 6.88
mm long. Palpi pale brown. Mandible ivory, teeth
fuscous. Clypeus convex, punctured, with a wide
polished margin. Face closely punctured, with grey
pilosity. Frons somewhat concave, transversely
wrinkled. Inner orbits with conspicuous yellow
margin. Outer orbits with narrow yellow margin
up to vertex. Malar space wide, with microsculp-
ture. Gena polished, with minute punctation, im-
plantations of grey pilosity. Tyloids on flagellar
segments 17-20. Prothorax with coarse sculpture
and robust epomia. Mesoscutum with rather fine
punctation, notaulus strong. Postscutellum with
yellow mark. Propodeum, with coarse sculpture,
both transverse carinae present, the apical carina
with small apophyses. Mesopleurum with coarse
punctation. All coxae black. Front and middle
femora robust and brownish in colour. Front and
middle tibiae brown and with a yellow streak. Hind
femur, tibia and basitarsus brown, third and fourth
tarsal segments paler. Entire gaster orange to red-
dish, claspers black.
There are two specimens from two islands, viz.,
Palma and Gomera. The position of the tyloids is
colour colour abdomen colour
femur 3 mandible
fuscous all tergites bright orange ivory; teeth
tip somewhat fuscous fuscous
black postpet. terg. 2, 3, 4,5 ivory; teeth
orange fuscous
6 & 7 somewhat
fuscous
slightly different from the holotype of Tenerife.
The other characters are worth to accept.
Male, paratype. Palma, La Rosa. Front wing 5.77
mm long. Base of mandible ivory. Face with pilos-
ity (spoilt). Sculpture of frons agreeing with holo-
type. Inner and outer orbits in part with ivory
margin. Tyloids on flagellar segments 16-19. Co-
lour of legs and abdomen agreeing with holotype.
Male, paratype. Gomera, Laguna Grande. Front
wing 6.66 mm long. Ivory spot on upper margin of
mandible. Face with grey pilosity. Sculpture of
frons agreeing with holotype. Inner and outer or-
bits in part with ivory margin. Tyloids on flagellar
segments 17-19 (16 minute) left antenna; 16-19
right antenna. Colour of legs and abdomen corres-
ponding with holotype. Compare table 4.
No additional material found. I did not see the
Canary Islands female. The characters given in the
key are based on the continental female of Cryptus
obscuripes.
Etymology. - “Impeditus” is the Latin for “dif-
ficult to approach”, relating to the difficulty to
place the specimens taxonomically.
The main characters of the male of C. obscuripes
and the subspecies smpeditus are shown on the
tables 3 and 4.
Cryptus nigritarsis Kriechbaumer
Cryptus nigritarsis Kriechbaumer, 1894: 45-46.
Cryptus nigritarsis; Ortega & Baez 1980: 49-50.
Material examined. — Lectotype by present designa-
tion: Santa Cruz, Teneriffa, 19.ii.92, Dr. Brauns leg. /
Cryptus nigritarsus 6, m. [handwriting of Kriechbau-
mer] / Lectotype label of present author. Paralectotype:
Santa Cruz, Teneriffa, 19.11.1892, leg. Dr. Brauns /
Itamoplex sp. 14 [label of Townes 1971] / Paralectotype
label of present author. Both types in Transvaal Museum,
Pretoria.
Other material: Two males: Tenerife, P. del Hidalgo,
2.v.1971, leg. Teunissen (coll. Van Rossem). Male. Tene-
rife, Las Cafiodas, 5.vi.1970 ? (date illegible), leg. G.
Ortega (Museo Insular Ciencias Nat. HY 1349).
VAN ROSSEM: Cryptus of Canary Islands 255
Table 4. Cryptus obscuripes impeditus subsp. nov.
locality tyloids colour inner colour outer
orbits orbits
Tenerife 17-20 ivory marked upper orbit
Liii.66 ivory
Palma 16-19 ivory marked ivory marked
4.viii.81
Gomera 1 17-19 ivory marked ivory marked
16.viii.77 r 16-19
Description of male lectotype. - Front wing 8.0
mm long. Palpi black. Base of mandible with yellow
spot. Labrum with white spots. Base of mandible,
gena, temple, frons and face with long silvery hairs.
Clypeus with a broad polished front part. Face
closely punctured to coriaceous towards eye mar-
gin. Frons coarsely wrinkled. Inner orbits with yel-
low margin. Temporal orbits with yellow marking.
Antenna with tyloids on flagellar segments 14-18.
Antennal scrobes developed. Pronotum coarsely
wrinkled, epomia present, black in colour with
ivory spoton upper margin opposite notaulus. Me-
soscutum with rough sculpture on front part of
median lobe and transverse wrinkling along notau-
lus. Lateral lobe with strong punctures. Notaulus
well developed. Scutellum with widely placed
rough punctures. Propodeum with basal and apical
transverse carina present, coarsely wrinkled. A pi-
cal carina with very small apophyses. Mesopleu-
rum with rough and regular sculpture. All coxae
and trochanters black. Front and middle tibiae and
femora orange in colour, tarsi brown. Hind femur
fuscous, laterally reddish brown. Hind tibiae and
tarsi black. First abdominal segment black, margin
of postpetiole orange. Other segments bright
orange. All tergites polished, with microsculpture.
Male claspers black. |
Description of male paralectotype. - Tyloids on
flagellar segments 14-18. Petiole and postpetiole
black, margin of postpetiole red. Other tergites
orange-reddish in colour. Male claspers black. Co-
lour of hind femur fuscous, laterally somewhat fer-
ruginous.
The main characters of C. nigritarsis are shown
on Table 5.
Cryptus praefortis spec. nov.
Material examined. - Holotype female: Tenerife,
Ijuana, 11.vii.1986, leg. G. Ortega / Museo Insular Cien-
colour colour abdomen colour
femur 3 mandible
brown entirely orange-red; yellow;
clasper black teeth fuscous
black entirely orange 1vory;
claspers black teeth fuscous
black entirely orange ivory spot
claspers black on upper margin
cias Nat. HY 6842 / Holotype label of present author.
Paratype female: Gomera (Isl. Can.), Llanos Crispin,
13.viii.1977, leg. M. Baéz / Museo Ins. Cienc. Nat. HY
1374.
Two female specimens from the islands of Tene-
rife and Gomera respectively, belong to an undes-
cribed species of the genus Cryptus showing affin-
ity with C. spinosus Gravenhorst and C. subspi-
nosus Smits van Burgst. The difference with
C. spinosus lies in the coriaceous sculpture of the
tergites in C. spinosus while in C. subspinosus the
ovipositor is longer, 0.50-0.60 of the length of the
front wing and the second, third and fourth tergites
show a close and fine alutaceous sculpture.
Description of female holotype. - Front wing
6.35 mm long. Palpi, mandible and entire head
black, except for minute yellow margins of inner
and outer orbits. Clypeus convex, with a flattened
front margin. Face coriaceous, medially with a cir-
cular protuberance. Lower frons rather strongly
concave, transversely wrinkled. Antenna black,
postanellus 5.6 X as long as apically wide. Malar
space wide. Pronotum coarsely wrinkled, epomia
strong. Mesoscutum polished, widely punctured,
notaulus strong and long, transversely wrinkled
along the sides of the median lobe. Scutellum pol-
ished, with fine punctures. Propodeum with coarse
sculpture, both transverse carinae present, the ap-
ical carina with strong apophyses, reminding of
C. subspinosus and C. spinosus. Mesopleurum
strongly sculptured. All coxae black and all other
parts of legs fuscous. The abdomen shows a main
character of the species, namely all tergites highly
polished and red in colour. Ovipositor 0.44 of the
length of the front wing.
The paratype closely resembles the holotype.
The ovipositor measures 0.47 of the length of the
front wing.
256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Table 5. Cryptus nigritarsis Kriechbaumer
locality tyloids colour inner colour outer colour colour abdomen colour
orbits orbits femur 3 mandible
lectotype Tenerife 14-18 yellow line tempor. orbit fuscous; petiole black base with
Santa Cruz yellow lateral. tergites orange yellow spot
reddish
paratype Tenerife 14-18 fuscous; first tergite black, other
Santa Cruz lateral. tergites orange
ferruginous
Tenerife P. del 14-18 yellow line ‚tempor. orbit fuscous; tip first tergite black; black
Hid. 2.v.1977 yellow somewhat other tergites reddish
ferruginous
Tenerife P. del 14-17 yellow line tempor. orbit fuscous first tergite black, terg. black
Hid. 2.v.1977 yellow ferruginous 2 & 3 red, tip black
Tenerife las 14-17 yellow spot at black fuscous first tergite black, terg. 2 black
Cañodas anten. base & 3 red, others ferruginous
Etymology. — “Praefortis” is Latin for “very black, apical part of postpetiole red. All other ter-
brave”. gites red coloured. Tergites subpolished, with mi-
Cryptus ultramondanus spec. nov.
Material examined. - Holotype female: Lanzarote (Isl.
Can.), Yaiza, 19.11.1979, leg. W. Perraudin (coll. Horst-
mann, Würzburg) / Holotype label of the present author.
Paratypes: Same data as holotype, 1 &; Lanzarote, Los
Valles, 20.11.1979, leg. M. Baéz, 1 ® (Museo Insul. Cienc.,
Nat HY 1327), 1 & (idem, HY 1346).
Description of female holotype. - Front wing
7.33 mm long. Palpi and mandible black. Clypeus
sparcely punctured, with a wide polished margin.
Face closely punctured. Malar space wide. Frons
wrinkled in front of anterior ocellus. Antennal
scrobes polished and somewhat concave. Inner or-
bit with yellow mark next to antennal socket.
Upper outer orbit with a small yellow spot. Gena
polished. Postanellus slender, 7.6 X as long as ap-
ically wide. Antenna with segments 5-9 white co-
loured. Pronotum coarsely punctured. Epomia
present. Mesoscutum and scutellum regularly
punctured. Notaulus strong. Mesopleurum with
coarse sculpture. Propodeum also coarsely sculp-
tured. Anterior transverse carina obsolete. The ap-
ical carina present, with weakly developed apo-
physes. All coxae black. Front femur and tibia and
middle tibia brownish in colour. Middle femur and
hind leg fuscous. Tarsi of all legs conspicuously
slender. Nervulus proximad of basal vein. Petiole
crosculpture. Ovipositor 0.86 of the length of the
front wing.
Description of the male paratype from Yaiza. —
The male agrees with the female. Not all characters
are recapitulated. Front wing 7.11 mm long. The
entire head, including the mandible, with long grey
setae. Right antenna missing beyond second flagel-
lar segment. Tyloids on flagellar segments 12-17.
Anterior transverse carina absent. The apical car-
ina with weakly developed apophyses. Margin of
postpetiole and all tergites orange in colour. Male
claspers black. Left hind leg lacking behind tro-
chanter.
Remark. - The male shows suspect resemblance
with the male of C. nigritarsis. The most impor-
tant difference lies in the colour of the front and
middle femora and tibiae, but it should be admitted,
in specimens from different islands. A definite de-
cision about conformity of the two species can only
be taken after the female of C. nigritarsis turns up.
Etymology. - “Ultramondanus” is Latin for “on
the other side of the world”.
The main characters of the male are shown on
tabel 6.
Cryptus insularis spec. nov.
Material examined. - Holotype male: Isl. Can., Fuertev
(= Fuerteventura), Vallebrin, 7.iii.1984, leg. G. Ortega /
VAN ROSSEM: Cryptus of Canary Islands Zo
Table 6. Cryptus ultramondanus spec. nov.
locality tyloids colour inner colour outer colour colour abdomen colour
orbits orbits femur 3 mandible
Lanzarote Yaiza 12-17 yellow spot yellow margin black first tergite fuscous; other black
19.11.79
at ant. scrobes temp. orbit
tergites red
Museo Ins. Cienc. Nat. HY 2162 / Holotype label of the
present author. Paratype: Same data, no. HY 2163.
Description of male holotype. - Front wing 5.06
mm long. Palpi and mandible fuscous. The entire
head black, only a minute marking of ivory on inner
orbit next to antennal socket and on temporal or-
bit. Face with close small punctures, implantations
of grey pilosity. Antennal scrobes polished and
somewhat concave. Upper frons not concave (com-
pare C. obscuripes), with weak sculpture around
anterior ocellus. OOL region, vertex and gena pol-
ished. Tyloids on flagellar segments 15-19. Lower
0.6 part of prothorax with wrinkled sculpture. Epo-
mia present. There is a vague light spot opposite
the notaulus. Mesoscutum polished with inconsid-
erable punctation. Scutellum polished. Basal area
and first lateral area, areola and second lateral area
polished (compare C. obscuripes). Both transverse
carinae present. Petiolar area wrinkled. Mesopleu-
rum with widely placed small punctures. All coxae,
including middle femur and entire hind leg black.
Front femur somewhat brownish. Front and mid-
dle tibia brownish, with an ivory streak. Entire
abdomen reddish, except for fuscous seventh ter-
gite and claspers.
The paratype agrees with the holotype. Tyloids
on flagellar segments 16-19. Orbits not marked.
Fifth, sixth and seventh tergites fuscous.
Etymology. - “Insularis” is Latin for “belonging
to the island”.
The main characters of the male are shown on
table 7.
Cryptus furvus spec. nov.
Material examined. - Holotype female: Gran Canaria,
Caldera Tejecta, Las Palmas, La Roche, 2.iv.1983 / Museo
Ins. Cienc. Nat. no. HY 1483 / Holotype label of the
present author.
Only one specimen of this species is available.
Within the genus Cryptus it takes an exceptional
position with its long ovipositor, 1.13 of the length
of the front wing.
Description of female holotype. - Front wing
7.33 mm long. Head including the mandibles black,
except for slight yellow marking on inner orbit
next to antennal socket. There is also a spot on the
outer orbit. Face with close punctation and coriace-
ous sculpture. Antennal scrobes polished, some-
what concave. Frons with coarse wrinkled sculp-
ture. Gena with widely placed fine punctures,
implantations of grey pilosity. Antenna long and
slender, sixth, seventh and eighth flagellar seg-
ments marked white. Pronotum with coarse sculp-
ture, the epomia robust. Lateral lobes of mesoscu-
tum polished, with weak punctures. The median
lobe proximally more closely punctured. The no-
taulus strong, with close transverse striation
across. Propodeum with coarsely wrinkled sculp-
ture. The basal transverse carina obsolete, the ap-
ical transverse carina present with weak apo-
physes. Mesopleurum coarsely sculptured. Legs,
including all coxae, fuscous. The front tibia slightly
swollen. The entire gaster black, with all tergites
highly polished, hairs almost obsolete. The highly
polished tergites offer a principal character. The
very long ovipositor, 1.13 of the length of the front
wing is another main character.
Etymology. - “Furvus” is the Latin for “dressed
in black”, relating to the entire black colour of the
specimen.
Cryptus verutus Van Rossem
Cryptus verutus Van Rossem, 1971: 209-211.
Cryptus verutus; Ortega & Baez 1980: 53-55.
Material examined. — Isl. Can. Tenerife, Güimar,
3.v.1965, leg. I. Klimesh, female. Holotype Van Rossem
(Naturhistorisches Museum, Wien), Isl. Can., Hierro,
9.iv.1898, leg. Hinz V, Female. Holotype of C. verutus
hierroensis Van Rossem (Museum für Naturkunde, Ber-
lin, DDR). Female, Tenerife, Tahodio, 25.11.1951, leg. R.
Arozena (Museo Ins. Cienc. Nat. no. HY 1357).
Description of female. - Front wing 7.5 mm
long. Head black with ivory lining of inner and
outer orbits. Postanellus 6.0-7.4 X as long as ap-
ically lining of inner and outer orbits. Postanellus
6.0-7.4 X as long as apically wide. Frons rugosely
wrinkled. Thorax roughly sculptured. Propodeum
258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Table 7. Cryptus insularis spec. nov.
locality tyloids colour inner colour outer colour colour abdomen colour
orbits orbits femur 3 mandible
holotype 15-19 marked yellow minute yell. black tergite 1-6 orange-red base light
Fuerteventura 1984 next ant.scrob. spot temp.orb. terg. 7 fuscous coloured
paratype 16-19 minute yellow black black tergites 1-4 red; vague light
Fuerteventura 1984 spot vertex
with anterior transverse carina obsolete. Posterior
transverse carina present with weak apophyses. All
femora orange. Abdomen with petiole fuscous, all
other tergites orange coloured. Ovipositor long,
0.96-1.0 of the length of the front wing.
Male unknown to me.
REFERENCES
Gravenhorst, J. L. C., 1829. Ichneumonologia Europaea
2(2): 1-989. - Vratislaviae.
Horstmann, K., 1968. Typenrevision der von Zetterstedt
beschriebenen Ichneumonidenarten. - Opuscula En-
tomologica 33: 305-323.
Kriechbaumer, J., 1894. Hymenoptera ichneumonidea. —
Berliner Entomologische Zeitschrift 39: 43-68.
Ortega, G. & M. Baéz, 1980. Contribucion al conocimiento
de los Ichneumonidos de las Islas Canarias. I. Subfami-
lia Gelinae (Hym. Ichneum.). - Annuario de Estudios
Atlanticos 26: 15-107.
other terg. fuscous spot at base
Rossem, G. van, 1969. A revision of the genus Cryptus
Fabricius s.str. in the western Palearctic region, with
keys to genera of Cryptina and species of Cryptus. —
Tijdschrift voor Entomologie 112: 299-274.
Rossem, G. van, 1971. A new species of Cryptus from the
Canary Islands. — Tijdschrift voor Entomologie 114:
209-211.
Rossem, G. van, 1987. Cryptus Fabricius, 1804 (Insecta,
Hymenoptera): proposed conservation. — Bulletin of
Zoological Nomenclature 44(1): 9-10.
Townes, H., 1970. The genera of Ichneumonidae part 2.
- Memoirs of the American Entomological Institute
We ESS
Zetterstedt, J. W., 1838. Insecta Lapponica 317-476. —
Lipsiae.
Received: 30 October 1988
Revised version accepted: 31 August 1989
A REVISION OF THE NEW WORLD GENUS NEURAESCHNA
HAGEN, 1867 (ODONATA: AESHNIDAE)
by
JEAN BELLE
ABSTRACT
Belle, J., 1989. A revision of the New World genus Newraeschna Hagen, 1867 (Odonata:
Aeshnidae). - Tijdschrift voor Entomologie 132: 259-284, figs. 1-75, tab. 1. [ISSN 0040-7496].
Published 1 December 1989.
A synopsis is given of the twelve representatives of the genus Neuraeschna Hagen. The
species are classified into groups. Separate keys to the males and females are constructed.
N. inarmata is synonymized with N. dentigera Martin, and N. rostrifera Martin is considered
to be a synonym of Heliaeschna simplicia (Karsch), a species confined to SE Asia. Five new
taxa are described and illustrated, viz. N. cornuta (male holotype: Suriname, Distr. Nickerie,
Sipaliwini), N. maya (male holotype: Costa Rica, Prov. Limón, Barra de Tortuguero),
N.mayoruna (female holotype: Peru, Dept. Loreto, Iquitos), N. maxima (male holotype: Brazil,
State of Para, Belem), and N. titania (male holotype: Ecuador, Prov. of Pichincha, Jaruqui).
Lectotypes are designated for three species described by Martin, viz. N. claviforctpata, N.
dentigera and N. harpya. The larva type of Newraeschna is determined by a reared individual
of N. harpya.
Key words. - South and Central America; Newraeschna; new species.
Dr. J. Belle, Onder de Beumkes 35, 6883 HC Velp, The Netherlands.
CONTENTS
IMEEOdUCHo NER EN 259)
Disposition of material studied and
ACKMOW IEG SE Me Mts yess O EN 260
SVSCEMAMESECHO DEFINITA boos aa cue 260
Treatment of the species
Key to the males of Neuraeschna............... 261
Key to the females of Neuraeschna ............. 262
Moe INE HOO 263
iNedenticeraiMartins 1000 ni. 263
N. mina Williamson & Williamson, 1930 ...... 264
NERC OMILULAES EE Neree nde Ce Sas CSN. 264
The Ni claviforcipata group... 265
N. claviforcipata Martin, 1909............... 265
(thepNeicostal:s croupeer ayes vst stai ae 266
Ni costalis (Burmeister, 1839)... 266
IN Sbroductaslimmins 9193.4) 222 22.2 20 een: 267
NMa vert nains OS nn eee ele ne 268
INL GLAD SPES OON ere 268
IN ZAND ESE DON ra een EE 269
MEN aaa IR EIN ERP ad oe 270
IN nb arMartin st 909s ee ones 270
DNErN 22724 BTOUPE 3: en. e Bae. 271
IN GOEETHOD. SSS (NON 2.2.2.8 SL Gede or 271
UDEINB AV Or knar CLOUD ta
IN|, GOGPORTLE SUG ON as oa ones sabes uo 2D
INEunaenchnaspetindet Mn ee Did
The status of Newraeschna rostrifera Martin, 1909 273
RIELELENCESN ERE ARO EEDE AEN RTE EN 2005
INTRODUCTION
The New World genus Newraeschna Hagen,
1867 includes at least twelve species of small to
very large Aeshnidae which are instantly recog-
nizable in the venation of the wing by the presence
of cross-veins in the midbasal space and in having
the subcosta prolonged through and beyond the
nodus.
The species of Neuraeschna are mainly Neo-
tropical in distribution but one specimen is here
recorded from the subtropical region of Brazil.
They haunt in woodlands. Their breeding places
seem mainly to be swampy areas but the only larva
of Newraeschna was taken in a bush creek. At
sunset the adults suddenly appear, sometimes in
troupes, to hunt in the dusk along the edges of
the woods and along the river banks near over
bush and water plants. Due to this crepuscular
habit and the fact that their flight is very erratic
and rather swift, these dragonflies are rarely col-
lected. As soon as the darkness falls more inten-
sively, they disappear again.
Davies and Tobin (1985) listed nine species of
Neuraeschna but two species should be discarded
since Newraeschna rostrifera Martin proves to fall
beyond the limits of the genus while Newraeschna
260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
inarmata Kimmins is found to be a junior synonym
of Newraeschna dentigera Martin. Here five more
taxa of Neuraeschna are introduced under the
names N. cornuta, N. maya, N. mayoruna, N. max-
ima and N. titania bringing the total number of
Newraeschna species to twelve.
Lectotypes are designated for the three Martin
species Newraeschna claviforcipata, N. dentigera
and N. harpya. These three species are redescribed
in common with the type-species N. costalis (Bur-
meister). The hitherto unknown larva-type of
Newraeschna could be determined by a reared in-
dividual of N. harpya.
An attempt is undertaken to divide the genus
Neuraeschna into species-groups. Separate keys
for the males and females have been prepared.
Unless otherwise indicated, the illustrations are
camera lucida drawings made by me (details com-
pleted by free-hand). The photographs have been
made by the museum photographers of the Rijks-
museum van Natuurlijke Historie, Leiden, and the
British Museum (Natural History), London. The
reproductions of these photographs are at three-
quarters of the full size of the dragonflies. The
Comstock-Needham terminology of the wing-
veins is used.
DISPOSITON OF MATERIAL STUDIED AND
ACKNOWLEDGEMENTS
The material on which the present review is
based, belongs to the institutions and personal col-
lections listed below; the names are preceded by the
acronyms used throughout the text of this paper;
they are followed by the names of the persons who
made this material accessible for this study. These
persons are most gratefully acknowledged here.
ANSP - Academy of Natural Sciences, Philadelphia;
Mr. Donald Azuma.
BMNH - British Museum (Natural History), London;
Mr. D. E. Kimmins (Ÿ) and Mr. Stephen J.
Brooks.
CG — Collection Garrison, Azusa; Dr. Rosser W.
Garrison.
CH — Collection Hellebuyck, Sherbrooke; Mr. Vic-
tor Hellebuyck.
CM — Collection Machado, Belo Horizonte; Prof.
Dr. Angelo B. M. Machado.
FSCA — Florida State Collection of Arthropods, Gai-
nesville; Prof. Dr. Minter J. Westfall, Jr.
IRSN - Institut Royal des Sciences Naturelles, Brus-
sels; Dr. Georges Demoulin and Dr. P. Groo-
taert.
MNHP - Muséum National d'Histoire Naturelle, Pa-
ris; Dr. Jean Legrand.
NHMV - Naturhistorisches Museum, Vienna; Dr. A.
Kaltenbach.
RNHL - Rijksmuseum van Natuurlijke Historie,
Leiden; Mr. J. van Tol.
UCV - Instituto de Zoologia Agricola, Universidad
Central de Venezuela, Maracay; Mr. Jorge de
Marmels.
UMAA - University of Michigan, Ann Arbor; Mrs. L.
K. Gloyd.
USNM - National Museum of Natural History,
Smithsonian Institution, Washington, D. C.;
Dr. Oliver S. Flint, Jr.
ZMHB - Zoologisches Museum, Humboldt Universi-
tit, East Berlin; Dr. Kurt K. Günther.
SYSTEMATIC SECTION
The name Newraeschna was proposed by Selys
but the genus was firstly characterized by Hagen
(1867: 54) and therefore it must be credited to
Hagen. Hagen compared Nezraeschna with its
closest relative Staurophlebia Brauer which has
also the subcosta prolonged through and beyond
the nodus but the midbasal space of Stawrophlebia
is free from cross-veins.
In addition to the characteristics given antea in
the introduction, the species of Newraeschna are
characterized by the following features:
Head. Superior surface of frons generally dark
brown in anterior part and pale brown in basal part
but one species has a well-marked brown T-spot.
Rear of head light brown with a narrow black
stripe bordering posterior margin of eyes from
occipital triangle to lateral emargination in eye
border.
Pterothorax. Predominantly brown with pale
markings as follows: On each lateral side of dorsum
a pale antehumeral marking representing a more
or less ovoid spot prolonged below in a narrow
point at lateral end of transverse mesepisternal
ridge or representing a more or less stripe-shaped
marking which tapers below and which generally
abruptly widens at upper edge. Lateral sides of
pterothorax with two complete pale stripes paral-
lel to the sutures and more or less of even breadth
throughout, one pale stripe on the mesepimeron
and the other on the metepimeron. Posterior to
these pale stripes the two sclerites are often black-
ish brown to black. Metepisternum with a more or
less subtriangular pale dorsal spot, and there is a
small pale spot close above the spiracle.
Wings. Contrary to Staurophlebia the costal and
subcostal interspaces posses accessory cross-veins
proximal to the first primary antenodal. Some spe-
cies, however, lack the basal costal cross-veins and
have only basal subcostal cross-veins. A marked
brown costal stripe is often present, especially in
the larger species. Membranule well-developed
and pale dirty white. Male anal triangle in hind
BELLE:
wing three-celled but there is sometimes an inter-
polated cell.
Abdomen of male. Swollen at base and con-
stricted on segment 3, then, in dorsal view, grad-
ually widening to rear of segment 8, 9 or 10. Dor-
sum of segment 9 raised convex and generally
denticulated in basal half, depressed concave and
not denticulated on apical half. Accessory genitalia
of similar type to those of Staurophlebia. Base of
inferior anal appendage with a dorsal prolongation
that is notably bulbous in the larger species (fig.
34).
Abdomen of female. Also swollen at base but in
general moderately constricted on segment 3, then,
in dorsal view, almost parallel-sided or gradually
narrowing to segment seven. Ventral process on
tenth segment two-pronged and usually with sub-
sidiary teeth at anterior side of base of fork, the
subsidiary teeth often discernible in a caudal view
of the fork.
The larva of Newraeschna is easily distinguished
from that of Stazrophlebia by the marked protrud-
ing eyes, by the lack of knobs on head and thorax,
and by the lack of a comb-like external prominence
on either mandible.
Treatment of the species
There are excellent characters available in the
structure of the male anal appendages for separat-
ing species of Newraeschna. The accessory genitalia
are conversely of little importance since they show
no or hardly differences in closely allied species.
The tenth sternite of the female abdomen exhibits
characters which can be used sometimes for the
determination of species. The features of the tho-
racic colour pattern are kept off in the keys because
this pattern often disappear completely or almost
Revision of Neuraeschna 261
completely through post mortem discoloration and
this applies especially to the females. Table 1 is an
alphabetic list of all valid taxa. The genus
Neuraeschna is provisionally divided into six spe-
cies groups which show some overlapping.
Within each group the species are treated chro-
nologically. For each species is given a list of the
main references, the new material and a descrip-
tion or descriptive notes.
KEY TO THE MALES OF NEURAESCHNA
(The male of N. mayoruna is unknown)
1. Superior surface of frons with a well-marked
brown T-spot (fig. 14) (N. harpya group) ...
see, VOI RR OA N. harpya
— Superior surface of frons without well-marked
brown T-spot but largely or anteriorly dark
brown shading to pale light brown at base 2
2. Small species. Hind wing < 54 mm ....... 3
— Large species. Hind wing > 54 mm ....... 6
3. Superior appendages with a stout, submedian,
dentiform dorsal process. Inferior appendage
about three-fourths the length of superiors (N.
dentigera group)
— Superior appendages without such a dorsal
process. Inferior appendage half as long as
superiors (N. claviforcipata group) .........
a i Re tete N. claviforcipata
4. Dentiform process of superior appendage very
large, its base about one-third the length of
appendage. Inferior appendage in profile view
nearly straight for its proximal two-thirds,
than curved slightly dorsad and again straight
in\itsidistal:third\(fig. 53,54) N. mina
- Dentiform process much smaller. Inferior ap-
pendage in profile view bent throughout its
wholéilengehi eine weet ae: 5
Table 1. - Alphabetic list of names of the species of Newraeschna, with type locality, type status and type location.
Species Type locality Type
holo lecto sex location
1. calverti Kimmins, 1951 Peru (eastern) X é BMNH
2. claviforcipata Martin, 1909 “Amazon region” X é IRSN
3. cornuta spec. nov. Sipaliwini, Surinam X é RNHL
4. costalis (Burmeister, 1839) Bahia, Brazil X Q NHMV
5. dentigera Martin, 1909 Turuty, Brazil x 8 IRSN
6. harpya Martin, 1909 “Amazon region” X é IRSN
7. maya spec. nov. Barra de Tortuguero, X 6) FSCA
Costa Rica
8. mayoruna spec. nov. Iquitos, Peru x Q UMAA
9. maxima spec. nov. Belem, Brazil X 8 RNHL
10. mina Williamson & Williamson, 1930 Porto Velho, Brazil X é UMAA
11. producta Kimmins, 1933 Iquitos, Peru x é BMNH
12. titania spec. nov. Jaruqui, Ecuador X é BMNH
262
10.
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
. Superior appendages with a truncated apex.
Dorsum of base of inferior appendage inflated
and discernible in dorsal view of abdomen (fig.
ALDO TAR AUTON RA AY RE N. dentigera
Apex of superior appendages tapering off to a
point. Dorsum of base of inferior appendage
notantlated! (figs DD) Wer ae ARC N. cornuta
. Superior appendages more or less flattened
and widened; in dorsal view the outer margin
of each appendage almost straight (N. costalis
group)
Superior appendages long, slender and forci-
pate; in dorsal view the outer margin of each
appendage strongly convex from base to apex
(fig. 57) (N. maxima group) .... N. maxima
. Superior appendage in profile view with the
upper edge of the raised apical portion pro-
duced in a long robust point (figs. 36, 43) ...
AAN DE OER SHES, LIO N. costalis
Superior appendage in profile view with the
raised apical portion appearing more or less as
a rhomboidal projection with round angles
. First angulation of inner margin of superior
appendage with a peg-like or spine-like projec-
tion. (EHESS TAMARA EN... N. calverti
First angulation of inner margin of superior
appendage rounded and smooth
. Dorsal prolongation of base of inferior appen-
dage in profile view not projecting above su-
perior appendage. Superior appendage with
outer apical margin angled and sometimes
bearinpfatdistincHpo BS In aa. 10
Dorsal prolongation of base of inferior appen-
dage in profile view projecting above superior
appendage. Superior appendage with outer ap-
ical margin broadly rounded (figs. 48, 49) ...
SAREI ari N. producta
Outer apical angle of superior appendage bear-
ing a small but distinct laterally directed sharp
point (fig. 51). Wings with well-developed
brown costal stripes, the brown colour of the
stripes extending over interspace behind RI,
between arculus and subnodus. Costal edge of
pterostigma in fore wing 5.5-5.8 mm, in hind
wing ACT N. titanta
Outer apical angle of superior appendage with-
out sharp point, at the most with a small tub-
ercle at inferior margin. Wings without or with
rather weakly developed brown costal stripes,
the brown colour of the stripes not extending
over interspace behind R1, between arculus
and subnodus. Costal edge of pterostigma in
fore wing 4.2-5.0 mm, in hind wing 3.2-4.0
ahaaa El dir reine ht N. maya
KEY TO THE FEMALES OF NEURAESCHNA
(The females of N. cornuta and N. mina are un-
known)
1.
Superior surface of frons with a well-marked
brown T-spot (fig. 14). Hind wing 55-60 mm
(N. harpya group) ............. N. harpya
Superior surface of frons without well-marked
brown T-spot but anteriorly pale or brown
shading to light brown at base ........... 2
. Small species. Hind wing < 54 mm ....... 3
Larger species. Hind wing > 54 mm ...... 4
. Wings with diffuse brown basal patches which
in antenodal interspaces are sometimes ex-
tended nearly to second primary antenodal.
Hind wing 50-51 mm; width of hind wing at
nodus 14 mm. Costal edge of pterostigma in
fore wing 40-45 mm (N. claviforcipata
group): 5.01. akk A N. claviforcipata
Wings without basal spots. Hind wing 46-48
mm; width of hind wing at nodus 13 mm.
Costal edge of pterostigma in fore wing 3.5-4.0
mm (N. dentigera group) … N. dentigera
. No basal costal cross-vein, only basal subcostal
cross-veins present. Fork of tenth sternite in
profile view distinctly rising before the line of
posterior margin of sternite (fig. 22). Hind
wing 61 mm; width of hind wing at nodus 17.5
mm. Costal edge of pterostigma in fore wing
4.5 mm (N. mayoruna group)
OR ES SORE N. mayoruna
Basal costal and subcostal cross-veins present.
Fork of tenth sternite in profile view not or not
notably rising before the line of posterior mar-
gin of sternite
. Abdomen swollen on segments 1 and 2, then
strongly constricted on segment 3; in dorsal (or
ventral) view the abdomen is three times as
wide at segment 2 as it is in the middle of
segment 3 (fig. 59). Hind wing 70 mm; width
of hind wing at nodus 19 mm. Costal edge of
pterostigma in fore wing 6 mm (N. maxima
SLOUP)L ee er o PEEETE N. maxima
Abdomen also swollen on segments 1 and 2 but
moderately or slightly constricted on segment
3 (N. costalis group)
. Superior surface of frons largely pale and with
a weakly developed brown central spot (fig.
15). Hind wing 66 mm; width of hind wing at
nodus 18.5 mm. Costal edge of pterostigma in
fore wing 6.3 mm. Specimen from subtropical
region of southern Brazil .... N. spec. indet.
Superior surface of frons with a brown anterior
marking extending over the whole breadth of
the frons
BELLE:
7. Vertex brown. Hind wing 59 mm; width of
hind wing at nodus 16.5 mm. Costal edge of
pterostigma in fore wing 5 mm. Wings with-
out or with weakly developed brown costal
SCRPEM TE Oil N. calverti
- Vertex black. Hind wing larger and broader 8
8. Fork of tenth sternite in profile view strongly
angled caudad, the prongs notably slender and
long, and curved at two-thirds length (figs. 18,
26). Hind wing 65 mm; width of hind wing at
nodus 17-18.5 mm. Costal edge of pterostigma
Im KORE vs DIO NATI O AI
a al en sr) N. producta
- Fork of tenth sternite not or not so strongly
curved caudad, the prongs not as in the preced-
na spese a LOR LE e 9
9. Large species with relative short pterostigma.
Hind wing 66-68 mm; width of hind wing at
nodus 18-19.5 mm. Costal edge of pterostigma
in fore wing 4.9-5.4 mm (= 7.7%-8.1% of
fore wing length). Central American spe-
CICERO. N. maya
- Pterostigma relatively longer, that of fore wing
9%-10% of fore wing length .......... 10
10. Large species with well-developed brown cos-
tal stripes, the brown colour of the costal stripe
extending over adjoining interspace behind
RI, between arculus and subnodus (figs. 72,
73). Tenth sternite robust, the fork in profile
view generally continuing the line of posterior
margin of sternite (figs. 20, 28). Hind wing
63-68 mm; width of hind wing at nodus
18-19.5 mm. Costal edge of pterostigma in
LOI SOIN 529245 qatar N. titania
- Somewhat smaller species. The brown costal
stripes generally not so well-developed as in
the preceding species. Tenth sternite less ro-
bust, the fork in profile view generally angled
caudad (figs. 17, 24). Hind wing 59-64 mm;
width of hind wing at nodus 17-18 mm. Costal
edge of pterostigma in fore wing 5-6.3 mm
3 65515 NE EREA N. costalis
THE N. DENTIGERA GROUP
In this group I place N. dentigera, N. mina and
N. cornuta which are bound together by the follow-
ing characters: (1) small species (hind wing < 54
mm); (2) no basal costal cross-veins but at least one
basal subcostal cross-vein present; (3) Superior
surface of frons without well-marked brown T-
spot; (4) pale antehumeral marking stripe-shaped
with pointed lower end and generally widened
upper end: (5) male superior appendage blade-
shaped and provided with a stout dentiform pro-
cess at or slightly before mid-length.
Revision of Neuraeschna 263
Species of the N. dentigera group: N. dentigera
Martin, N. mina Williamson & Williamson and N.
cornuta spec. nov.
Neuraeschna dentigera Martin, 1909
(figs. 1, 41, 42, 64)
Neuraeschna dentigera Martin, 1909: 207, 208, fig. 213
(4 app.) - 69 Surinam; Williamson & Williamson
1930: 15 (4, ® Guyana); Kimmins 1951: 48.
Neuraeschna inarmata Kimmins, 1951: 46-48, figs. 8,9
(8 app.). - @ New River, Guyana. Syn. nov.
Material. - Brazil: state of Para; Rio Xingu Camp, ca.
60 km S. of Altamira (52° 22’ W., 3° 39’ S.), 8 October
1986, 1 9 (Igarape, mist nets at dusk); 9 October 1986,
1 2 (Ist jungle stream, trail 1); 9 October 1986, 1 9 (mist
net at dusk), all P. Spangler & O. Flint, USNM. - Guyana:
Essequibo River, Rockstone, 1 February 1912, 1 3, 1 9,
L. A. & E. B. Williamson & B. J. Rainey, UMAA. - Peru:
Dept. Loreto, Prov. Maynas, Iquitos, 8 July 1931, 1 8
(teneral), from Paul Nagel, FSCA.
Lectotype designation of Neuraeschna dentigera
Martin, 1909. - According to Martin (1909) the
original series of Newraeschna dentigera is in the
Selysian collection and consists of two males and
two females from “Surinam”. However, I found
three males and one female while, from the pin
labels, these specimens appeared to be secured in
Turuty at the Amazon River by Bates. The chosen
lectotype is the male which has served for fig. 213
in Martin's monograph of 1909. This male is dis-
tinguished from the other two of the original series
by the somewhat more strongly curved inferior
appendage. The pin labels are “31”, “Turuty”,
“Neuraeschna dentigera Bates @ Turuty” (in Selys’
hand; the words “modesta Selys” deleted and
changed into “dentigera Bates”), “Collection Selys
Type Neuraeschna dentigera Bates Revision Mar-
tin 190 Neuraeschna dentigera Martin” (text
partly printed) and “Type Neuraeschna dentigera
Martin” (the word “Type” printed in red). I have
added the yellow pin label “LECTOTYPE Rev. J.
Belle, 1988”.
The lectotype is in fairly good condition but the
tips of the right pair of wings are broken off distal
to the pterostigmata.
At the end of the description of Neuraeschna
inarmata, Kimmins (1951) stated that his species
is closely related to Neuraeschna dentigera Martin
but readily separable from it by the form of the
male superior anal appendage “unless his (= Mar-
tin’s) figure is very inaccurate”. Because fig. 213 in
Martin's monograph shows the superior anal ap-
pendage with a second tooth-like projection on the
264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
inner margin of each appendage in contrast to
Kimmin’s male which has not such an extra (se-
cond) tooth.
On 19 April 1961 I visited the Brussels Museum
in order to check fig. 213 in Martin's monograph.
The depiction proved to be very inaccurate, indeed.
In stead of a second tooth-like projection there was
a broad rounded lobe. Also the inferior appendage
appeared not so strongly curved as depicted. At my
visit of the British Museum (Natural History) on
20 June 1961 I was able to compare the males of
Neuraeschna dentigera from the Selysian collec-
tion with the holotype male of Newraeschna inar-
mata. No character was found that would justify a
specific distinctness of the two species. Thus we
consider Nezraeschna inarmata Kimmins a junior
synonym of Newraeschna dentigera Martin.
Male. - Total length 64-69 mm; abdomen 49-53
mm (incl. app. 5.0-5.2 mm); hind wing 43-46 mm;
costal edge of pterostigma in fore wing 3.9-4.0
mm, in hind wing 2.9-3.0 mm.
Head. Face orange-brown, the anteclypeus with
greyish tinge. Superior surface of frons dark brown
anteriorly, fading to pale light brown basally. Ver-
tex dark brown. Antennae light brown. Occipital
triangle brown. Upper part of rear of head black.
Pterothorax: Dark brown with pale (= green)
stripes. Pale antehumeral markings stripe-shaped,
the upper part widened mesad, the lower third
tapering to lateral end of transverse mesepisternal
ridge (fig. 1). Pale mesepimeral and metepimeral
stripes about equal in width. A small pale dorsal
spot on metepisternum. Wing articulations with
pale spots.
Legs. Reddish brown but dark brown at knees.
Wings. Clear, in old specimens faintly brown
tinged. Pterostigma light brown. Membranule
dirty white, that of hind wing extending along
posterior wing margin to a point about two-fifths
the way along first paranal cell. Antenodals in fore
wing 23-28, in hind wing 19-21.
Abdomen. Dark brown but segment 2 with
green middorsal stripe. In dorsal view constricted
on segment 3, then gradually wider on segments 4
to 9 with the segments 6 to 9 becoming more
depressed successively. Upper surface of segment
10 flat and elevated at apex. Appendages dark
brown and shaped as shown in figs. 41 and 42.
Female. — Total length 65-67 mm (excl. app);
abdomen 50-54 mm (excl. app.); hind wing 46-48
mm; width of hind wing at nodus 12.5-13.5 mm;
costal edge of pterostigma in fore wing 4.0-5.0
mm, in hind wing 2.7-3.0 mm.
Similar to male regarding stature and general
coloration but dark brown colour of pterostigma
less dark and abdominal segment 2 not constricted.
Antenodals in fore wing 23-30, in hind wing
19-22. Fork of tenth sternite in profile view con-
tinuing the line of posterior margin of sternite.
Anterior surface of sternite at base of fork with two
or three small subsidiary teeth on either side.
Save the deviations in the curving of the inferior
appendage the males also differ in the develop-
ment of the tooth-like projection and the confor-
mation of the apical portions of the superior ap-
pendages. These have or have not a small acute
tooth on the outer apical end.
Neuraeschna mina Williamson & Williamson,
1930
(fig. 53, 54)
Neuraeschna mina Williamson & Williamson, 1930:
9-15, figs. 1, 2 (& app.) - 6, Brazil (Territorio de
Rondônia).
Neuraeschna cornuta spec. nov.
(figs. 2, 55, 56)
Material. - Holotype: Suriname: Distr. Nickerie, Sipa-
liwini (at a small creek along a patch to the airstrip), 14
February 1961, 1 g, D.C. Geijskes, RNHL.
Male (holotype; a pinned specimen in perfect
condition). — Total length 67 mm; abdomen 51.5
mm (incl. app. 4.9 mm); hind wing 44 mm; costal
edge of pterostigma in fore wing 3.3 mm, in hind
wing 2.5 mm.
Head. Face light brown but upper half of vertical
part of frons dark brown. Superior surface of frons
dark brown on anterior half with convex posterior
margin (no T-spot), the basal half pale light brown.
Antennae light brown. Vertex black. Occipital tri-
angle dark brown.
Pterothorax. Dark brown with pale (= green)
markings. Antehumeral markings stripe-shaped,
the lower end tapering and directed to lateral edge
of anterior mesepisternal ridge, the upper end
slightly widened mesad but much widened laterad
(fig. 2). Pale mesepimeral stripe rather narrow.
Pale metepimeral stripe as wide as pale mesepi-
meral stripe at its upper end, becoming gradually
narrower toward below.
Wings. With a faint brown tinge, the costal mar-
gins slightly darker. Pterostigma brown-yellow.
Membranule reaching to a point about one-third
the way along first paranal cell. Antenodals in fore
wings 28-30, in hind wings 22-23.
Abdomen. Dark brown with pale (= yellow to
BELLE:
yellow-green) markings on segments 2 and 3. Seg-
ment 2 with a broad pale streak along ventral tergal
margins and behind auricles; middorsum of seg-
ment with a pale stripe over whole length of seg-
ment, not connected with the pair of two short,
transverse pale stripes at posterior border of seg-
ment. Segment 3 with large pale basal side spots
extending along ventral tergal margin to halfway
the segment, and with a pair of small, transversely
elongated, dorsal posterior pale spots. Segment 3
constricted. Apical segments depressed. Anal ap-
pendages shaped as shown in figs. 55 and 56, the
superiors being a trifle longer than segments 9 and
10 together.
THE N. CLAVIFORCIPATA GROUP
N. claviforcipata shows the features listed for
the N. dentigera group except for point (6). In N.
claviforcipata the dorsal dentiform process is re-
placed by a mere raised outer edge of the superior
appendage on the basal half.
Species of the N. claviforcipata group: N. clavı-
forcipata Martin.
Neuraeschna claviforcipata Martin, 1909
(figs. 3, 4, 16, 33, 39, 40, 65)
Neuraeschna claviforcipata Martin, 1909: 206, 207, fig.
212 (8 app.) - 6 ® Amazon region; Williamson &
Williamson 1930: 15 (8 Brazil, State of Pará); Racenis
1970: 29 (4 Venezuela); Geijskes 1971: 664 (4 French
Guiana).
Material. — Brazil: State of Amazonas, Rio Negro (at
Manaus), 2 8, ex coll. Lacroix, MNHP; State of Pará, 1
6; State of Pará, June 1935, 1 &, G. V. Vredenburg,
BMNH; State of Pará, Belem, 5 August 1922, 1 & (ten-
eral), J. H. Williamson, UMAA. - Ecuador: Prov. Napo,
Rio Napo, Limoncocha (300 m), 5 November 1980, 1 9
(teneral), M. J. Westfall, Jr. & David Robinson, FSCA. —
Suriname: Distr. Marowijne, Tapanahoni River, Drietab-
betje (in forest), 8 October 1952, 1 9, D. C. Geijskes,
RNHL. - Venezuela: Est. Bolivar, Guayaraca-Auyante-
qui, 16 April 1956, 1 8, J. Racenis, UCV.
Lectotype designation of Newraeschna clavifor-
cipata Martin, 1909. — For this purpose the best
preserved male of the original series of this species
in the Selysian collection (IRSN) has been taken.
The pin labels are “109”, “Bates”, “Neuraeschna
claviforceps Bates & Amazone” (in Selys’ hand),
“Collection Selys Type Neuraeschna claviforcipata
Bates Revision Martin 190 Neuraeschna clavifor-
cipata Martin” (text partly printed) and ‘Type
Neuraeschna claviforcipata Martin” (the word
“Type” printed in red). I have added the yellow pin
label “LECTOTYPE Rev. J. Belle, 1988”.
Revision of Neuraeschna 265
The wings of the lectotype are not perfect. The
tip of the right fore wing is broken off proximal to
the pterostigma while the hind border of the hind
wings displays some damages.
Male. - Total length 70-75 mm; abdomen 56-58
mm (incl. app. 4.5-5.0 mm); hind wing 49-51 mm;
costal edge of pterostigma in fore wing 3.2-4.0
mm, in hind wing 2.5-3.0 mm.
Head. Face light brown. Superior surface of
frons dark brown anteriorly, fading to pale light
brown basally. Antennae brown, the first antennal
segment darker. Vertex black. Occipital triangle
black.
Pterothorax. Dark brown with pale stripes. An-
tehumeral markings green to bluish-green, stripe-
shaped, the upper part widened mesad, the lower
third tapering to lateral end of transverse epister-
nal ridge (figs. 3, 4). A moderately wide green
stripe On mesepimeron and an equal one on mete-
pimeron. A small green dorsal spot on metepister-
num. Wing articulations green.
Legs. Red-brown, darker at knees and on tarsi
and claws. >
Wings. Clear, in aged individuals slightly brown
tinged with faintly discernible brown costal stripes.
Pterostigma light brown. Membranule of hind
wing extending along posterior margin of wing to
a point nearly halfway along first paranal cell. An-
tenodals in fore wing 27-36, in hind wing 19-25.
Abdomen. Long and slender. Segment 3 con-
stricted. Segments 7 to 10 distinctly depressed.
Dark brown with pale (= yellow to yellow-green)
markings on segments 1 to 4. Segment 1 with two
connected pale dorsolateral spots. Segment 2 pale
behind auricles and along ventral tergal margins,
with a fine pale middorsal stripe from base to a
point three-fourths the way along segment, and
with a submedian dorsal pair and a more posterior
dorsal pair of transversely elongated pale spots.
Dorsum of segment 3 with two small pale spots at
transverse carinae and another pair of small pale
spots at posterior margin. Dorsum of segment 4
with two small pale spots near posterior margin.
Appendages dark brown, about as long as segments
9 and 10 together, shaped as shown in figs. 39 and
40.
Female. - Total length 67-69 mm (excl. app.);
abdomen 51-54 mm (excl. app.); hind wing 50-51
mm; width of hind wing at nodus 14 mm; costal
edge of pterostigma in fore wing 3.8-4.5 mm, in
hind wing 2.8-3.1 mm.
Similar to male but abdominal segment 3 not
constricted and end segments of abdomen not de-
pressed. Wings clear with diffuse brown basal
266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
patches which in antenodal interspaces are some-
times extended to second primary antenodal (fig.
65). Antenodals in fore wing 29-32, in hind wing
21-23. Fork of sternite of abdominal segment 10
shaped as shown in fig. 16.
The measurements here mentioned are those of
fully mature specimens. They are smaller in the
teneral specimens. The teneral male from Belem
has the dimensions: total length 64 mm, abdomen
50 mm (incl. app. 4.5 mm) and hind wing 46 mm;
those of the teneral female from Limoncocha are:
total length 64 mm (excl. app.), abdomen 50 mm
(excl. app. 5.5 mm) and hind wing 47 mm.
The teneral female from Limoncocha is the only
female with intact anal appendages. Each appen-
dage is slender and narrow on the basal third; the
apical two-thirds portion is blade-shaped and has
a rounded median keel on the upper surface (fig.
33). The fork of the tenth sternite has on its ante-
rior surface a relatively large central supplemen-
tary tooth which is discernible in caudal view.
There is not such a supplementary tooth in the
other females.
The prongs of the fork of the tenth sternite are
more divergent in the complete syntype female
than in the other females. The fork of the other
syntype female is broken away.
THE N. COSTALIS GROUP
The characters of this groupe are: (1) large spe-
cies (hind wing > 54 mm); (2) basal costal and
subcostal cross-veins present; (3) superior surface
of frons without well-marked brown T-spot; (4)
pale antehumeral marking more or less ovoid, the
lower end pointed, the upper end sometimes
slightly constricted; (5) male superior appendages
blade-shaped without any kind of a dorsal process
on basal half.
The members composing the N. costalis group:
N. costalis (Burmeister), N. producta Kimmins, N.
calverti Kimmins, N. titania spec. nov. and N.
maya spec. nov.
Neuraeschna costalis (Burmeister, 1839)
(figs. 5, 17, 24, 31, 34, 35, 36, 43, 66-68)
Aeschna costalis Burmeister, 1839: 837 — Q Brazil, State
of Bahia.
Gynacantha ferox Erichson, 1848: 585 (& Guiana, holo-
type in ZMHB); Hagen 1867: 54.
Neuraeschna costalis, Hagen 1867: 55, 56; Selys 1883:
748 (40 sep.); Kirby 1890: 95; Martin 1909: 205, fig.
210 (8 app.); Kimmins 1933: 226; Räcenis 1970: 29,
30 (& Venezuela); Geijskes 1971: 663 (9 French Gui-
ana); Davies & Tobin 1985: 15.
Material. - Brazil: State of Pará, Obidos, 1 @, ANSP (ex
coll. Calvert); Obidos, March 1971, 1 4, B. Ferreira; Dis-
trito Federal, Brasilia, May 1970, 1 9, Frits, CM. State of
S20 Paulo, Rio Guaratuba (Varjào), 7 March 1969, 3 4,
1 2; 29-30 April 1969, 9 6, 5 9; 21 April, 2 6, 1 9; 23
April 1970 (at 6 p.m.), 1 9; 15 April 1971 (at 5.30 p.m.;
riverbank), 1 &; 19 April 1971 (at 5.30 p.m.; riverbank),
2 &; 19 March 1972, 2 9, all E. Dente, CM but 2 4, 1 9
in RNHL. - Ecuador, Prov. Pastaza, Rio Napo, Puyo-
Oriente (elev. 1000 m), 29 November 1936 (at 5.30 p.m.),
1 @, W. Clark-Macintyre, UMAA. - Guyana: Mazaruni-
Potario Dist, Kartabo Point; Earthwatch Research
Exped. 21 December 1983, 1 9, W. E. Steiner, USNM. —
Suriname: Distr. Suriname, Paramaribo (Cultuurtuin), 9
November 1938, 1 9; Distr. Saramaca, Coesewijne River
(at light), 25 October 1945, 1 Q; Distr. Marowijne, be-
tween Moengotapoe and Wia Wia (trail 19, at swamp),
October 1948, 2 4; Marowijne River (first island), 23
October 1948, 1 @, all D. C. Geijskes; Distr. Saramacca,
Garnizoenspad (km 20, at 6.30 p.m.), 7 September 1955,
2 4; Distr. Suriname, Fernandesweg naar Zee, 9 Sep-
tember 1955, 1 &, 1 9; Para River (along road to Dom-
burg, at 6.15 p.m.), 21 September 1955, 1 &, Rijsdijkweg,
4 March 1957, 1 4, Paramaribo (Zorg en Hoop), 19
August 1958, 1 9; 17 August 1959, 1 9, all J. Belle; Distr.
Marowijne, Albina, August 1973, 1 @ (teneral); 28 Sep-
tember 1973, 1 9; Distr. Suriname, Zanderij, 13 Sep-
tember 1973, 1 Q, all J. J. Belle, RNHL. Distr. Suriname,
Paramaribo (Zorg en Hoop), 22 August 1958, 1 9; Zan-
derij (Weg naar Matta), 28 July 1963, 1 8; both J. Belle,
ZMHB. - Venezuela: 1899, 1 4, F. Gray, MHNP. Est.
Amazonas, Simarawochi, 30 March 1973, 1 6, W. Perez
(No. 13892). Est. Bolivar, Uruyen, 11 April 1956, 1 6,
Foldat (No. JR-06637); El Bochinche, 5-8 December
1985, 1 9, L. D. Otero & A. Chacón, UCV.
Male. - Total length 82-91 mm; abdomen 62-71
mm (incl. app. 5.9-6.5 mm); hind wing 56-62 mm;
costal edge of pterostigma in fore wing 5.0-5.8
mm, in hind wing 3.6-4.6 mm.
Head. Face yellow-brown to brown but antecly-
peus with a tinge of green. Frons granulous. Super-
ior surface of frons dark brown anteriorly, fading
to pale light brown basally; the posterior margin of
the dark brown marking straight or convex. Vertex
dark brown to black. Antennae light brown. Occip-
ital triangle dark brown and tufted with brown
hairs. Eyes in freshly killed specimens dark green
on upper half and yellow-green on lower half.
Pterothorax. Dark brown with pale markings.
Antehumeral pale markings blue-green, ovoid, its
antero-lateral corner prolonged in a narrowed
point reaching to lateral end of transverse mesep-
isternal ridge (fig. 5). The two pale lateral stripes
rather narrow and green, followed by a dark brown
to black stripe. Wing articulations dark blue.
Legs. Red-brown, darker at knees.
Wings. Clear, the brown costal band variable, in
some specimens developed only at base of wings,
BELLE:
in other specimens well-developed and extending
over adjoining interspaces behind R1. In aged in-
dividuals the wings becoming dark brown tinged,
often firstly in the middle from discoidal triangle
to pterostigma. Venation dark brown. Pterostigma
brownish yellow. Membranule dirty white, that of
hind wing extending to a point halfway along first
paranal cell. Antenodals in fore wing 31-36, in
hind 23-26.
Abdomen: Dark brown. Constricted on segment
3, becoming gradually wider and more depressed
on segments 4 to 10. Anal appendages shaped as
shown in figs. 35 and 36.
Female. - Total length 81-85 mm (excl. app);
abdomen 61-64 mm (excl. app. 3.5-4.0 mm); hind
wing 59-64 mm; width of hind wing at nodus
17-18 mm; costal edge of pterostigma in fore wing
5.0-6.2 mm, in hind wing 4.0-5.3 mm.
Coloration similar to male. Abdomen slightly
constricted on segment 3, then in dorsal view al-
most parallel-sided on segments 4 to 7, slightly
wider on segments 8 and 9. Fork of tenth sternite
in profile view generally distinctly angled caudad
(fig. 17) and with one to four subsidiary teeth on
anterior basal side of fork. Anal appendages an-
tenna-shaped and short, about as long as abdomi-
nal segment 9 (fig. 31). Antenodals in fore wing
30-37, in hind wing 23-27.
Infraspecific variation. - The specimens from
Suriname are the smallest of the series, those from
southern Brazil the largest.
The male anal appendages show some interest-
ing morphological variations. In some males the
raised apical portion of the superior appendage in
profile view appears as a rhomboidal projection
with an upper apical point but in other males more
or less as a triangular projection (fig. 43). The male
from Obidos in the Machado collection has the
upper apical points of the superiors notably con-
verging; they are generally parallel or diverging.
The inferior appendage shows variation in its cur-
vature and in the degree of swollenness of its
basal dorsal prolongation. The tip of the inte-
rior appendage reaches to a point between two-
thirds and four-fifths of the way along the super-
iors as the base of the inferior may be more or less
hidden between the superiors and the anal tuber-
cles.
The thoracic colour pattern is variable and espe-
cially the pale antehumeral markings can consider-
ably vary in size. The males from Venezuela have
a relatively small pale antehumeral spot in the
centre of each lateral side of the thoracic dorsum.
In other specimens the pale antehumeral marking
Revision of Neuraeschna 267
occupies almost the whole lateral side of the tho-
racic dorsum.
The specimens of Newraeschna exhibit also a
typical colour-heteromorphism with regard to the
brown costal stripes. I distinguish:
Type. This is of course the form in the holotype
(fig. 66). The brown colour of the stripe is not
extended over the adjoining interspace behind R1,
between arculus and subnodus.
Var. hyalinata nov. Here the brown costal stripe
is absent or reduced to a mere diffuse brown basal
patch. Specimens pertaining to var. hyalinata seem
only to occur in the Guianas and Venezuela (fig. 67,
68).
Var. marginata nov. Specimens of this variation
have definite brown costal stripes with the brown
colour of the stripes extended over the adjoining
interspace behind R1, between arculus and subno-
dus. In the material before me the two males from
Bolivar, Venezuela and the female from Brasilia,
Brazil belong to var. marginata. The brown costal
stripes of these examples approach in develop-
ment those of N. titania spec. nov. (figs. 72, 73).
Neuraeschna producta Kimmins, 1933
(figs. 6, 18, 26, 48, 49)
Neuraeschna producta Kimmins, 1933: 226-228, figs. 1,
2 (6 app.) - 4 9 Peru; 1951: 45, 46, figs. 5-7 (¢ hw-
base & tip of app., fork of 9 10th sternite).
Material. - Brazil: State of Pará. Tapajos River (Bar-
reira), 1 g, IRSN.
One of the male paratypes (dated 25 September
1930) ex coll. Hincks in the Manchester Museum
has been carried over to the Rijksmuseum van
Natuurlijke Historie at Leiden. This male has
weakly developed brown costal stripes with the
brown colour of the stripe not extended over the
interspace behind R1, between arculus and subno-
dus. The present male from Barreira, however, has
hyaline wings with well-developed brown costal
stripes, the brown of the stripe being extended
over the interspace behind R1, between arculus and
subnodus.
The female allotype is notably larger than the
males, has relatively broader wings and well-deve-
loped brown costal stripes with the brown colour
of the stripe extended over the interspace behind
R1, between arculus and subnodus. The colour
pattern of the pterothorax is lost through post
mortem discoloration. The hind margin of the
brown anterior marking on the superior surface of
the frons is strongly convex in the female allotype;
it is straight in the males. In the Selysian collection
268 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
there is an Amazonian female collected by Bates
which possibly belongs to N. producta. This fe-
male, with relatively narrower wings (hind wing
65 mm; width of hind wing at nodus 17.5 mm), has
the posterior margin of the brown anterior mark-
ing of the frons straight. Unfortunately the fork of
the tenth sternite is broken away and due to this
damage the specimen cannot well be compared
with the female allotype.
Neuraeschna calverti Kimmins, 1951
(figs. 7, 8, 19, 27, 45-47, 70, 71)
Neuraeschna calverti Kimmins, 1951: 43-46, figs. 1-4 (8
app. & hw base, fork of 9 10th sternite) - & Q Peru.
Material. - Brazil: State of Amazonas, Manaus, no date,
1 g, Vieira leg., CM. - Suriname: Distr. Suriname, Zan-
derij, 18 December 1973, 1 4, J. J. Belle, RNHL.
Two illustrations in the original description of
N. calverti are inaccurate. In the figure of the male
appendages in dorsal view (Kimmins’ fig. 1) the
dorsal prolongation of the lamina supra-analis is
not depicted while in the figure of the fork of the
tenth sternite of the female (Kimmins' fig. 4) the
subsidiary teeth are missing. The present figures
47 and 19 are improved versions of these figures.
Only three males of N. calverti are known. The
differences noticed between them and listed below
seem to me not of specific value.
(1) Peg-like projection on inner margin of su-
perior anal appendage small and very acutely
pointed in the male from Suriname (fig. 45).
(2) Inner apical projection of superior appen-
dage longer in the male from Manaus (fig. 46).
(3) Pterostigma different in size; costal edge of
pterostigma of fore wing 4.5 mm in the male holo-
type, 5.0 mm in the male from Manaus, and 5.5 mm
in the male from Suriname.
(4) Pale antehumeral marking smallest and very
similar to that of the males of N. costalis from
Venezuela in the male from Suriname (fig. 7); the
pale antehumeral marking is largest and almost
completely occupying the lateral half of the tho-
racic dorsum in the male from Manaus (fig. 8).
(5) Anterior lamina of accessory genitalia
stouter in the male from Manaus.
Neuraeschna titania spec. nov.
(figs. 9, 20, 28, 50-52, 72, 73)
Material. - Holotype: Ecuador, Prov. Pichincha, Jaru-
qui, 1 & (from Staudinger) in BMNH. Paratypes: Ecua-
dor: Prov. Napo, Archidona (675 m), May 1977 (at light),
1 9, Herman G. Real, CG; Prov. Pastaza, Puyo (wa-
tershed Rio Pastaza-Rio Napo, 1000 m), 21 November
1936 (at twilight), 1 ® (teneral); 28 November 1936, 1
Q (allotype); watershed Rio Arajuno-Río Napo (head-
waters, 1000 m), 29 April 1941, 1 9, all W. Clark-Macin-
tyre; Prov. Zamora-Chinchipe, Amora (700 m), 3 De-
cember 1941, 1 @ David B. Laddey, UMAA; Prov.
Pinchincha, Jaruqui, ,1 9, from Staudinger, BMNH —
Peru: Prov. Lima, vicinity of San Pedro (900 m), 26 May
1935, 1 @ (flying at dusk), Felix Woytkowski, UMAA.
Male (holotype; an aged individual; tip of right
fore wing broken away; fig. 72). - Total length 90
mm; abdomen 70 mm (incl. app. 5.5 mm); hind
wing 64 mm; costal edge of pterostigma in fore
wing 5.5 mm, in hind wing 4.5 mm.
Head. Face largely pale brownish yellow shading
to pale brown above. Superior surface of frons
anteriorly dark brown shading to pale light brown
basally, the posterior margin of the dark brown
margin strongly convex and reaching nearly to
base. Vertex black. Antennae brown. Occipital tri-
angle brown, provided with brown hairs.
Pterothorax. Dark chocolate brown, the brown
colour darkest immediately posterior to the two
greenish yellow lateral stripes. On each lateral side
of the thoracic dorsum a large, bluish green ovoid
spot prolonged below in a narrowed point at lateral
end of transverse mesepisternal ridge (fig. 9). The
two pale lateral stripes slender and about equal in
width.
Legs. Third femur brown, second femur darker
brown, first femur blackish brown with brown-
yellow inner side. Tibiae reddish brown. Knees and
tarsi darker brown.
Wings. Brown tinged with a marked brown cos-
tal stripe, the brown colour of the costal stripe
extending over adjoining interspace behind RI,
between arculus and subnodus. Venation dark
brown but costa pale brown anteriorly. Ptero-
stigma brown-yellow. Membranule extending
along hind wing margin toa point half way the first
paranal cell. Antenodals in fore wings 36-37, in
hind wings 25-30.
Abdomen. Dark brown with some indefinite
pale areas on lower part of sides of segments 1 and
2 and basal three-fourths of segment 3. Broad at
base. Segment 3 moderately constricted. Segments
4 to 9 gradually widening. Segments 7 to 9 gradu-
ally more depressed. Dorsum of segment 9 strongly
convex and densely denticulated in basal half, de-
pressed concave and not denticulated on apical half.
Anal appendages shaped as shown in figs. 51 and
52, the superiors a little shorter than segments 9
and 10 together.
Female (allotype; an aged individual; anal ap-
pendages broken away; tip of fore wings broken off
but not lost). - Total length 90 mm (excl. app.);
BELLE:
abdomen 67 mm (excl. app.); hind wing 68 mm;
greatest width of hind wing 19.5 mm; costal edge
of pterostigma in fore wing 6.4 mm, in hind wing
5.9 mm.
Similar to male holotype but abdomen slightly
constricted on segment 3, then gradually narrow-
ing to apex of segment 7. Fork of tenth sternite
continuing the line of the hind margin of the ster-
nite. Base of fork with a relatively large supple-
mentary tooth each side and well-developed minor
teeth on anterior surface (fig. 20, 28). Antenodals
in fore wings 38-39, in hind wings 27.
The male from Peru and the female from Ar-
chidona (Ecuador) are younger than the type spec-
imen. Their wings are hyaline and have definite
brown costal stripes. The male from Peru is also
the smallest male of the series (abdomen 67 mm;
hind wing 60 mm) and has a paler thoracic colour
pattern. Especially the anterior streaks of the two
pale lateral stripes are notably lighter. Like in the
males of the other species of the N. costalis group
the raised apical portion (comb) of the superior
appendages varies in height and length.
The apices of the anal appendages of the female
allotype are, like in the other females, broken away;
the remaining pieces are parallel-sided, 2 mm long
and 0.6 mm wide. The anal appendages are highly
probable antenna-shaped, like in N. costalis and N.
harpya. The fork of the tenth sternite of the female
paratype in the British Museum (Natural History)
is damaged, the right prong being broken away.
Neuraeschna maya spec. nov.
(figs. 10, 21, 25, 44, 74)
Material. — Holotype: Costa Rica: Prov. Limón, Barra
de Tortuguero, 17 July 1985, 1 & (Victor Hellebuyck) in
FSCA. Paratypes: Costa Rica: Prov. Limón, Tortuguero,
17 July 1985, 1 g; Barra de Tortuguero, 17 July 1985, 2
6, 1 (allotype); 18 July 1985, 6 6; 19 July 1985, 2 3,
all Victor Hellebuyck, CH, but 1 @ in RNHL. - Honduras:
1 9, IRSN; Puerto Cortez, 30 May 1917, 1 9, E J. Dyer,
USNM.
Male (holotype; an aged individual; head broken
off but not lost; distalia of antennae lost). - Total
length 88 mm; abdomen 67 mm (incl. app. 5.8
mm); hind wing 59 mm; costal edge of pterostigma
in fore wing 4.5 mm, in hind wing 3.5 mm.
Head. Face dull olive green. Dark anterior mark-
ing of superior surface of frons developed, its pos-
terior margin convex in middle. Base of superior
surface of frons pale light brown. Vertex black.
Scape of antenna black, pedicel yellowish brown.
Occipital triangle yellowish brown anteriorly, dark
brown posteriorly.
Revision of Neuraeschna 269
Pterothorax. Blackish brown white pale mark-
ings. On each lateral side of thoracic dorsum a
bluish-green ovoid spot which is not distinctly pro-
longed below on lateral side (fig. 10). Meso-para-
ptera (antealar sinus) largely yellow-green. Mes-
and metepimeral pale stripes rather narrow and
yellow-green.
Legs. Third femur brown, black at knees. Second
and first femora very dark brown. Tibiae brown,
tarsi and claws darker brown.
Wings. Brown tinged with faint brown costal
stripe. Venation dark brown but costae light brown
anteriorly. Pterostigma light brown. Membranule
ratherbroadandalong posterior wing margin reach-
ing to a point nearly halfway the first paranal cell.
Antenodals in fore wings 32-33, in hind wings 23.
Abdomen. Dark brown with some indefinite
paler areas on lower part of sides of segments 1 and
2 and base of segment 3. Broad on basal segments
1 and 2, constricted on segment 3, almost parallel
from apex of segment 3 to base of segment 7, than
gradually widening to apex of segment 8 and more
depressed to-segment 9. Segments 9 and 10 equal
in width. Dorsum of segment 9 strongly convex
and denticulated on basal half, depressed concave
and not denticulated on apical half. Anal appen-
dages very resembling those of N. tztanza but outer
apical angle of each superior appendage without a
sharp point. Instead there is a small tubercle on
inferior margin at the outer apical angle of the
superior appendages (fig. 44).
Female (allotype; a very aged individual; apices
of anal appendages broken away). - Total length 87
mm (excl. app.); abdomen 68 mm (excl. app.); hind
wing 65 mm; width of hind wing at nodus 18 mm;
costal edge of pterostigma in fore wing 4.9 mm, in
hind wing 3.8 mm.
Similar to male but abdomen not constricted on
segment 3 and then gradually narrowing to apex of
segment 7. Fork of sternite of tenth abdominal
segment angled caudad; anterior surface of base of
fork with two small subsidiary teeth on left side
only (fig. 21). Wings very dark brown tinged with
weakly discernible brown costal stripes. Anteno-
dals in fore wing 36, in hind wing 26.
All Costa Rican specimens are aged to very aged
individuals and due to their brown tinged wings the
brown costal stripes are not well discernible. The
two females from Honduras, however, are young
specimens (one teneral). The wings of these fe-
males are hyaline and the brown costal stripes can
well be studied (fig. 74). The brown colour of the
costal stripe is not extended over the interspace
behind R1, between arculus and subnodus.
270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
THE N. HARPYA GROUP
Although evidently related to the members of
the N. costalis group, N. barpya is aberrant from
its members and from all other species of the genus
by the presence of a well-marked brown T-spot on
the top of the frons. With a hind wing length of
52-60 mm, N. harpya belongs to the moderately
large species of the genus.
It is classified here as a separate group: N. har-
pya Martin.
Neuraeschna harpya Martin, 1909
(figs. 11, 14, 32, 37, 38, 60-63, 69)
Neuraeschna harpya Martin, 1909: 206, fig. 211 (6 app.)
— & 2 Amazon region, Suriname; Williamson & Wil-
liamson 1930: 15 (Q Guiana); Kimmins 1933: 226,
227; 1951: 48; Geijskes 1971: (4 Q French Guiana).
Material. - Brazil: State of Para, Tapajós (53 27), 1 6,
BMNH; State of Para, Rio Xingu Camp (52° 22’ W 3°
39’ S, ca. 60 km S. of Altamira, 1 October 1986 (at dusk),
1 Q; same camp, Igarape (N. of camp, trail 4), 9 October
1986, 1 4, P. Spangler & O. Flint, USNM; same camp and
collectors, 3 October 1986, 1 4, CG. Rio Negro, 20 August
1951, 1 Q, Pe. J. Falco; Rio Uaupés (Taracua), 14 August
1964, 1 ®, CM. - French Guiana: Camopi (on light), 19
November 1969, 1 g, Balachowski & Grüner, CM. —
Guiana: Essequibo River, Rockstone, 1 February 1912, 1
Q LA. & E.B. Williamson & B. J. Rainey, UMAA. - Peru:
“Dept. Loreto, Rio Ampi Yaco, Estiron, 29 March-9 April
1970, 1 © (teneral), B. Malkin, RNHL. - Surinam: 1 6,
H. Fruhstorfer (99-103); Distr. Marowijne, Litani River
(Feti Creek), 17 August 1939, 1 9, D. C. Geijskes, BMNH.
Distr. Marowijne, Litani River (Feti Creek ), 16-22 July
1939, 2 & 5 2. Distr. Suriname: Coropina Creek, Repu-
bliek, 27 May 1951, 2 g (teneral), all D. C. Geijskes. Distr.
Nickerie, Corantijn River, Wonotobo Falls, 1 February
1956, 1 & 1 2; Kabalebo River, 26 August 1963, 1 9;
Distr. Suriname, Zanderij, Troelinde Creek (Bos Beheer),
20 July 1958, 1 9; Zanderij, Pontji Creek, Dauwdrop-
kamp, 2 September 1958, 1 4; Zanderij, Pontjibrug, 19
November 1958, 1 9, all J. Belle; Distr. Nickerie, Wilhel-
mina Mountain Range, camp at Lucie River, 21 August
1963 (6.30 p.m.), 1 6, S. Ligorie; Corantijn River, Wono-
tobo Falls, 23 July 1959, 1 Q (reared), D. C. Geijskes,
RNHL.
Lectotype designation of Newraeschna harpya
Martin, 1909. - In the Selysian collection there are
three syntypes of Newraeschna harpya, two males
and one female. The male, which bears the pin
labels “Santarem”, “149 Æ. harpya B.” and “Collec-
tion Selys Neuraeschna harpya Bates @ Revision
Martin 190 Neuraeschna harpya Bates” (partly
printed), is here designated as the lectotype of
Neuraeschna harpya Martin, 1909. To the pin of
this specimen I have added the yellow label
“LECTOTYPE Rev. J. Belle, 1988”.
The lectotype is in fairly good condition. The
costae of the fore wings are broken just proximal
to the pterostigma but restored. The pterothorax
is broken between the second and third pair of legs
but otherwise all critical details are intact and can
well be studied.
Male. - Total length 73-74 mm; abdomen 57-58
mm (incl. app. 5 mm); hind wing 52-54 mm; costal
edge of pterostigma in fore wing 3.9-4.3 mm, in
hind wing 2.8-3.0 mm.
Head. Face yellow-brown. Superior surface of
frons with dark brown to black T-spot, the area
next to the stem of the T-spot raised convex and
pale light brown. Vertex black. Antennae brown.
Occipital triangle black. Eyes in freshly killed spec-
imens green above, becoming yellowish below and
bluish along black hind border.
Pterothorax. Brown with broad pale (= light
green) stripes. Pale antehumeral markings stripe-
shaped, the upper end widened on both sides, the
lower part becoming narrower and reaching lateral
end of transverse mesepimeral ridge (fig. 11). Pale
mes- and metepimeral stripes very broad and equal
in width, each pale lateral stripe posteriorly bor-
dered by a very dark brown stripe. A small green
dorsal spot on metepisternum. Wing articulations
with green to bluish green spots.
Legs. Dark brown, the inner side of first femur
green for the basal three-fourths portion.
Wings. Clear, sometimes with a faintly deve-
loped brown costal band. Venation dark brown but
frontal margin of costa lighter. Pterostigma light
brown. Antenodals in fore wing 28-36, in hind
wing 21-25. Membranule of hind wing extending
to a point one-third the way along first paranal cell.
Abdomen. Brown, the posterior margin of each
segment almost black. Segments 1 and 2 with light
green side spots. Segment 3 to 7 with a yellowish
green lateral spot at transverse carina, the one of
segment 3 extending on basal half of segment
along ventral tergal margin. Segment 3 somewhat
constricted, slightly wider on segments 4 to 9, suc-
cessively. Segment 10 narrower, its upper surface
flat and elevated at apex. Segments 6 to 9 becoming
more depressed, successively. Anal appendages
dark brown, shaped as shown in figs. 37 and 38.
Female. - Total length 73-78 mm (excl. app.);
abdomen 57-61 mm (excl. app. 3.5 mm), hind
wing 55-60 mm; costal edge of pterostigma in fore
wing 4.6-5.0 mm, in hind wing 3.0-3.5 mm.
Coloration similar to male. Abdomen very
slightly constricted on segment 3, in dorsal view
almost parallel-sided on segment 4 to 9, narrower
on segment 10. Anal appendage antenna-like, as
BELLE:
long as segment 9, in dorsal view almost parallel-
sided, slightly tapering at apex, the apical two-
fifths portion superiorly keeled (fig. 32). Fork of
tenth sternite in profile view almost continuing the
line of posterior margin of sternite, but sometimes
slightly curved caudad. There are often no subsi-
diary teeth on the anterior surface of the base of the
fork. Antenodals in fore wing 30-36, in hind wing
21-25.
Larva (described from a larval cast-off skin of a
reared female). - Total length 38.5 mm; abdomen
26.5 mm (incl. app. 3.7 mm); greatest width of
abdomen 8 mm; width of head over the eyes 9 mm;
third femur 7.5 mm.
Larval skin clean and practically hairless. It is
light brown, almost without colour pattern but
there are conspicuous dark markings on the
femora.
Head. Flat above. Eyes very prominent. Poste-
rior portion of occiput produced laterally with tap-
ering sides. Antennae 7-jointed, the first two basal
segments short and rounded, the third segment
longest and the fourth segment shortest (fig. 62).
Labium brown, its basal hinge reaching to third
coxae. Front margin of mentum with a pair of
rather long spines, each spine slightly curved out-
ward and located in the middle of the part between
the small median cleft and the base of the lateral
lobe. End hook of lateral lobe terminating in a
strong acute spine, its inner border nearly semicir-
cular. Movable hook strong, two and a half times
as long as outer border of lateral lobe (fig. 61).
Prothorax. Flat above. Dorsal disc with promi-
nent rounded lateral margins. Supracoxal armature
with the first proces conical and twice as small as
the second one.
Synthorax. Without knobs. Tips of wing-cases
reaching to well upon segment 4.
Legs. Femora flattened, distal third black, prox-
imal two-thirds of inner and outer surface with a
row of small black roundish spots. Claws simple.
Abdomen. Base of segment 7 with a middorsal
pair of black dashes. Widest on segments 6 and 7.
No dorsal hooks. Lateral spines on segments 5 to
10, the ones on segment 5 minute but distinct, the
ones on segments 8 and 9 largest, the ones on
segment 10 small. Inferior anal appendages
slightly shorter than segments 9 and 10 together,
the superiors about two-thirds as long as inferiors,
the middorsal appendage a trifle shorter than in-
feriors (fig. 63). Gonapophyses reaching backward
to well beyond segment 9.
The larva of Nezraeschna harpya was found in
Revision of Neuraeschna ZI
a small bush creek near the Wonotobo Falls by Dr.
D. C. Geijskes. It was in the transformation period
climbing up a grass stem at 9 a.m. After it was
taken to the camp at the Wonotobo Falls, the imago
appeared in the following night (Dr. Geijskes
1959, personal communication).
THE N. MAXIMA GROUP
N. maxima is characterized in both sexes by the
strongly constricted abdomen on the third seg-
ment, and in the male sex by the very long, slender
and forcipate superior anal appendages. Otherwise
the species agrees with the characters of the N.
costalis group.
Species of the N. maxima group: N. maxima
spec. nov.
Neuraeschna maxima spec. nov.
(figs. 12, 57-59, 75)
Material. - Holotype: Brazil; State of Parà, Belem, Val-
de-Can airport (5.30 a.m.), 11 September 1963, 1 @ (B.
Malkin) in RNHL. Paratypes: Brazil: State of Pará, Be-
lem (Utinga Forest), 10 October 1954, 1 8 1 9 (allotype),
both Inacio, CM; Belem, December 1954, 1 &, ex coll.
Agriculture Station (D. C. Geijskes).
Male (holotype) - Total length 100 mm; abdo-
men 78 mm (incl. app. 9 mm); hind wing 67 mm;
costal edge of pterostigma in fore wing 6 mm, in
hind wing 4.5 mm.
Head. Face brown but anteclypeus darker and
vertical part of frons blackish brown. Superior sur-
face of frons blackish brown on anterior half, pale
brown on basal half. Vertex black-brown. Scape of
antenna black, the pedicel and distalia brown. Oc-
cipital triangle brown-yellow.
Pterothorax. Dark colour brown. On each lateral
side of dorsum a large ovoid bluish green spot
prolonged below in a narrow point at lateral end
of transverse mesepisternal ridge (fig. 12). Mes-
and metepimeral pale stripes greenish yellow, the
area anterior to each pale lateral stripe lighter
brown and the area posterior to each pale lateral
stripe very dark brown.
Legs. Femora dark brown. Second and third ti-
biae red-brown, first tibia dark red-brown. Tarsi
and claws blackish brown.
Wings. Hyaline with a distinct brown costal
stripe extending from base to pterostigma, the
brown colour of the costal stripe extending over
adjoining interspace behind R1, between arculus
and subnodus. Pterostigma, light yellow. Anteno-
dals in fore wings 39, in hind wings 28-30.
Abdomen. Swollen on segments 1 and 2,
272 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
strongly constricted on segment 3, then gradually
widening to apex of segment 10. Convex basal half
of segment 9 nearly devoid of denticles. Anal ap-
pendages shaped as shown in figs. 57 and 58. Dor-
sal prolongation of base of inferior appendage
moderately developed, not rising above superior
appendages and not discernible in dorsal view of
abdomen.
Female (allotype; much broken but details well-
preserved; apices of anal appendages broken
away). - Total length 97 mm (excl. app.); abdomen
75 mm (excl. app.); hind wing 70 mm; width of
hind wing at nodus 18 mm; costal edge of ptero-
stigma in fore wing 6.0 mm, in hind wing 4.8 mm.
Very similar to male regarding stature and col-
oration. Wings also very similar to those of holo-
type. Antenodals in fore wings 33-44, in hind
wings 30-31. Fork of sternite of abdominal seg-
ment 10 shaped as in N. costalis. Anterior surface
of base of fork with three (left side) and four (right
side) small subsidiary teeth.
THE N. MAYORUNA GROUP
The unique specimen of N. mayoruna is a female
which agrees with the characters of the N. costalis
group except for point (2). N. mayoruna has no
basal costal cross-veins, only basal subcostal cross-
veins. In this respect it agrees with the members
of the N. dentigera group and N. claviforcipata
group. Further the female is distinguished from all
other species of the genus in having the fork of the
sternite of the tenth abdominal segment distinctly
rising before the posterior side of the sternite (fig.
22).
Species of the N. mayoruna group: N. mayoruna
spec. nov.
Neuraeschna mayoruna spec. nov.
(Lie seno 223920)
Material. - Holotype: Peru: Dept. Loreto, Mishquiyacu
(15 km from Iquitos), 29 July 1931, 1 Q (from Paul Nagel)
in UMAA.
Female (holotype; apices of anal appendages
broken away; wings slightly damaged). — Total
length 85 mm (excl. app.); abdomen 65 mm (excl.
app.); hind wing 61 mm; width of hind wing at
nodus 17.5 mm; costal edge of pterostigma in fore
wing 4.7 mm, in hind wing 3.5 mm.
Head. Face reddish brown-yellow. Superior sur-
face of frons dark brown on anterior half, pale
brown on basal half (fig. 13). Vertex black. Anten-
nae with black scapes, the pedicels and distalia
brown. Occipital triangle black.
Pterothorax. Obscured through post mortem
discolouration but there is something like a pale
stripe discernible on mes- and metepimeron.
Legs. Femora reddish brown. Dark brown at
knees, on tibiae, tarsi and claws.
Wings. Brown tinged with a distinct brown cos-
tal stripe extending from base to pterostigma; the
brown colour of the stripe faintly developed in
adjoining interspace behind R1, between arculus
and subnodus. Pterostigma yellow. Antenodals in
fore wing 41-45, in hind wing 32-33. Wings with
only a single basal subcostal cross-vein. Area be-
tween Cul and Cu2 with two rows of cells for its
proximal one-fourth portion (in left hind wing
with an extra third cell) followed by one row of
cells; Cu2 sagged out on this proximal one-fourth
portion.
Abdomen. Very dark brown, moderately swollen
on segments | and 2, not constricted on segment
3, then gradually narrowing to apex of segment 7.
Fork of sternite of tenth segment shaped as shown
in figs. 22 and 29; anterior surface of base of fork
with a subsidiary tooth on each side.
Neuraeschna spec. indet.
(figs. 15, 23, 30)
Material. — Brazil: State of Santa Catarina, Joinville, 1
Q, Staudinger & Bang-Haas vend., BMNH (ex coll. Cow-
ley, No. 9994).
Female (frons above transversely crushed; api-
ces of anal appendages broken away; wings slightly
damaged). — Total length 86.5 mm (excl. app.);
abdomen 66.5 mm (excl. app.); hind wing 66 mm;
width of hind wing at nodus 18.5 mm; costal edge
of pterostigma in fore wing 6.3 mm; in hind wing
5.0 mm.
Head. Face brown-yellow. Superior surface of
frons predominantly pale brown, the central por-
tion becoming darker brown, the anterior border
broadly brown-yellow (fig. 15). Vertex laterally
brown-yellow, in middle darker brown. Scape of
antenna dark brown, pedicel brown-yellow, distalia
brown.
Pterothorax. Due to post mortem discoloration
there are no pale markings discernible on the dor-
sum. Posterior half of mes- and metepimeron
blackish brown; each of these dark streaks preceded
by brown areas (the usual pale lateral stripes being
apparently discoloured).
Legs. Femora brown, dark brown at knees. Se-
cond and third tibiae red-brown. Tarsi, claws and
first tibia dark to very dark red-brown.
Wings. Hyaline proximal to nodus, brown
BELLE:
tinged distal to nodus. There are distinct brown
costal stripes extending from base to pterostigma,
the brown colour of the stripe not extending over
adjoining interspace behind R1, between arculus
and subnodus. Pterostigma yellow. Antenodals in
fore wings 36-37, in hindwings 24-26.
Abdomen: Dark brown. Moderately swollen on
segments 1 and 2, then gradually narrowing to
apex of segment 7. Fork of tenth sternite shaped
as shown in figs. 23 and 30; anterior surface of base
of fork with six small subsidiary teeth.
This female is the only specimen of Newr-
aeschna reported from the subtropical region. It
highly probable belongs to a member of the N.
costalis group. Though very closely allied with N.
costalis it seems to be inconspecific with it unless
the subtropical representatives of this species have
a paler frons and larger proportions. By the ab-
sence of sufficient clear distinguishing characters I
have refrained from giving it a name.
THE STATUS OF NEURAESCHNA ROSTRIFERA
MARTIN
In his monograph of 1909 (pag. 208, 209; fig.
214), Martin described Newraeschna rostrifera on
the basis of a unique male from Suriname. The late
lamented Dr. D. C. Geijskes, who was most inter-
ested in the odonate fauna of Suriname, located and
studied (June 1966) the male holotype of
Neuraeschna rostrifera in René Martin's collec-
tion. On his return from Paris he communicated
me that the type fell beyond the limits of the genus
Neuraeschna simply because the subcosta was not
prolonged through and after the nodus.
In my opinion the holotype of N. rostrifera is no
more than a variation of Heliaeschna simplicia
(Karsch) syn. nov., an Old World species described
and figured in Martin's same monograph (pag.
160, 161; fig. 161). The reference to the locality
“Surinam” on the pin label of the holotype of N.
rostrifera may be an error pro Sumatra, an island
where H. simplicia occurs (cf. Lieftinck, 1954: 104,
105).
Revision of Neuraeschna 273
REFERENCES
Burmeister, F., 1839. Handbuch der Entomologie II (2),
Berlin: 805-862.
Davies, D. A. & P. Tobin, 1985. The dragonflies of the
world. A systematic list of the extant species of Odo-
nata. Vol. 2. Anisoptera. - Rapid Communications
Societas Internationalis Odonatologica (Supple-
ments) 5: XII + 151 pp.
Erichson, W. F, 1848. Reisen in Britisch-Guiana in den
Jahren 1840-1844 (3): 553-617 (R. Schomburgk, ed.)
J.J. Weber, Leipzig (Libellulinae: 583-586).
Geijskes, D. C., 1971. List of Odonata known from French
Guyana, mainly based on a collection brought together
by the mission of the “Muséum National d'Histoire
Naturelle”, Paris, (1) (2). - Annales de la Société
Entomologique de France (N.S.) 7 (3): 655-677.
Hagen, H. A. 1867. Notizen beim Studium von Brauer’s
Novara-Neuropteren. - Verhandlungen der Zoolo-
gisch-Botanischen Gesellschaft in Wien 17: 31-62 (1-
32 sept.).
Kimmins, D. E., 1933. A new species of Neuraeschna
(Odonata). - The Entomologist 66: 226-228.
Kimmins, D. E., 1951. Two new South American Aesh-
nidae (Odonata). - Entomological News 62: 43-48.
Kirby, W. F., 1890. A synonymic catalogue of Neuroptera
Odonata, or dragonflies. With an appendix of fossil
species. - Gurney & Jackson, London: IX + 202 pp.
Lieftinck, M. A., 1954. Handlist of Malaysian Odonata. A
catalogue of the dragonflies of the Malay Peninsula,
Sumatra, Java and Borneo, including the adjacent
small islands. - Treubia 22, Supplement: i-xiii, 1-202,
map.
Martin, R., 1908-1909. Aeschnines. - Collections Zoolo-
giques du Baron Edm. de Selys Longchamps. Cata-
logue systématique et descriptif. Fasc. 18-20. Hayez,
Bruxelles. 223 pp.
Racenis, J., 1970. Los Odonatos de la region del Auyan-
tepui y de la Sierra de Lema, en la Guayana Venezo-
lana. 2. Las familias Gomphidae, Aeshnidae y Cordu-
liidae. - Acta Biologica Venezuelica 7 (1): 23-39.
Selys Longchamps, E. de, 1883. Synopsis des Aeschnines.
Premiere partie: Classification. — Bulletin de l'Acadé-
mie Royale de Belgique (3) 5: 712-748 (3-40 sep.).
Williamson, E. B. & J. H., 1930. Two new Neotropical
Aeshnines (Odonata). - Occasional papers, Museum
of Zoology, University of Michigan 218: 1-15, pl. 1.
Received: 1 August 1989
Accepted: 30 August 1989
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 1-12. Diagram of colour pattern of left lateral side of thoracic dorsum of males in Neuraeschna: 1, dentigera
lectotype); 2, cornuta (holotype); 3, claviforcipata (lectotype); 4, claviforcipata (Venezuela); 5, costalis (Surinam);
6, producta (paratype); 7, calverti (Surinam); 8, calverti (Brazil); 9, titania (paratype, Peru); 10 maya (holotype); 11,
arpya (lectotype); 12, maxima (holotype).
13-15. Colour pattern of superior surface of frons in Neuraeschna: 13, mayoruna (holotype); 14 harpya (8
ıriname); 15 spec. indet.
BELLE: Revision of Neuraeschna 275
21
dok 50}
Fig. 16-23. Left profile view of ventral process on abdominal segment 10 of female in Newraeschna: 16, claviforcipata
(Ecuador); 17, costalis (Brazil); 18, producta (paratype); 19, calverti (allotype); 20, titania (allotype); 21, maya
(allotype); 22, mayoruna (holotype); 23, spec. indet.
Figs. 24-30. view of ventral process on abdominal segment 10 of female in Neuraeschna: 24, costalis (Brazil); 25, maya
(allotype); 26, producta (paratype); 27, calverti (allotype); 28, titania (allotype); 29, mayoruna (holotype); 30, spec.
indet.
Figs. 31-33. Dorsal view of left anal appendage of female in Newraeschna: 31, costalis (Suriname); 32, harpya
(Surinam); 33, claviforcipata (Ecuador. Basal one-tenth portion of distorted appendage not depicted).
Fig. 34. Left profile view of lamina supra-analis (inferior appendages) of male in Newraeschna costalis (Brazil).
276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 35-42. Improved versions of figs. 210-213 in Martin, 1909. Neuraeschna costalis, 6: 35, tenth abdominal segment
and anal appendages, dorsal; 36, the same, left profile. Newraeschna harpya, 6: 37, tenth abdominal segment and anal
appendages, dorsal; 38, the same, left profile. Neuraeschna claviforcipata, &: 39, tenth abdominal segment and anal
appendages, dorsal; 40, the same, left profile. Newraeschna dentigera, 8: 41, tenth abdominal segment and anal
appendages, dorsal; 42, the same, left profile.
Fig. 43. Newraeschna costalis, 6. Apex of left superior appendage, left profile (BMNH).
Figs. 44-46. Dorsal view of apex of left superior appendage of male in Nezraeschna: 44, maya (holotype); 45, calverti
(Suriname); 46, calverti (Brazil).
BELLE: Revision of Neuraeschna Pi
56
Fig. 47. Neuraeschna calverti, &. Anal appendages, dorsal (improved version of fig. 1 in Kimmins, 1951). Figs. 48,
49. Neuraeschna producta, & (after Kimmins, 1933): 48, tenth abdominal segment and anal appendages, dorsal; 49,
the same, left profile.
Figs. 50-52. Neuraeschna titania, 6 holotype: 50, accessory genitalia, ventral; 51, apical segments of abdomen with
anal appendages, dorsal; 52, the same, left profile. Figs. 53, 54. Neuraeschna mina, & holotype (after Williamson &
Williamson, 1930): 53, anal appendages, dorsal; 54, tenth abdominal segment and anal appendagesin left profile. Figs.
55, 56. Newraeschna cornuta, 8 holotype: 55, tenth abdominal segment and anal appendages, dorsal; 56, the same,
left profile.
278 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 57-59. Neuraeschna maxima: 57, tenth abdominal segment and anal appendages of @ paratype, dorsal; 58, the
same, left profile; 59, basal segments of abdomen of 9 allotype, ventral.
Figs. 60-63. Neuraeschna harpya, 2 exuviae: 60, head, dorsal; 61, mentum, ventral; 62, right antenna, dorsal; 63, apical
segments of abdomen with anal appendages, left profile.
BELLE: Revision of Neuraeschna 279)
Figs. 64-65.—64, Neuraeschna dentigera Martin, @ syntype; 65, Newraeschna claviforcipata Martin, 9 Suriname.
280 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 66-67 —66, Newraeschna costalis (Burmeister), 9 holotype; 67, Neuraeschna costalis (Burmeister), var. hyalinata
nov., 2 Suriname.
BELLE: Revision of Neuraeschna 281
Figs. 68-69.—68, Neuraeschna costalis (Burmeister), var hyalinata nov, 8 Suriname; 69, Neuraeschna harpya Martin,
& Suriname.
282 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 70-71.—70, Neuraeschna calverti Kimmins, & holotype; 71, Newraeschna calverti Kimmins, 9 allotype.
BELLE: Revision of Neuraeschna
Figs. 72-73.—72, Neuraeschna titania spec. nov., & holotype; 73, Neuraeschna titania spec. nov., Q
same locality as holotype.
paratype from
284 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 74-75.—74, Neuraeschna maya spec. nov., Q paratype, Honduras (IRSN); 75, Newraeschna maxima spec. nov.,
6 paratype (RNHL).
A NEW SAPRINUS FROM IRIAN JAYA (COLEOPTERA:
HISTERIDAE)
P. KANAAR
ABSTRACT
Kanaar, P., 1989. A new Saprinus from Irian Jaya (Coleoptera, Histeridae). — Tijdschrift
voor Entomologie 132: 285-287, figs. 1-5 [ISSN 0040—7496]. Published 1 December 1989.
Saprinus grandiclava sp.n. is described from Irian Jaya. It belongs to the primitive group
of Australian and Pacific Saprinus-species with non-fused paramera.
Key words. — Histeridae, new species, Irian Jaya, Oriental region.
P. Kanaar, Irenestraat 8, 2351 GL Leiderdorp, The Netherlands
INTRODUCTION
In 1973 the Rijksmuseum van Natuurlijke His-
torie in Leiden obtained the beetle collection of
the late P. H. van Doesburg Sr. This collection
contains predominantly beetles collected in the
former Dutch East Indies. Among the unidentified
Histeridae in this collection I found some spec-
imens of a new Saprinus-species from Hollandia
in the former Dutch New Guinea (present name:
Jayapura, Irian Jaya, Indonesia). The description
is as follows:
Saprinus grandiclava sp.n.
(figs. 1—5)
Description
Body outline (fig. 1) broadly oval, moderately
convex. Length of body (head, propygidium and
pygidium excluded): 4.0 mm., width of body: 3.3
mm. Colour brownish black, shiny; vertex and e-
lytra metallic blue, pronotum with cupreous me-
tallic lustre.
Head: Mandibulae regularly bent, convex, punc-
tured, tip sharply pointed. Labrum transverse,
emarginate. Clypeus continuous with front, hardly
convex, finely punctate. Front flat, moderately and
scarcely punctate. Frontal stria interrupted, con-
tinuous with supraorbicular striae. Eyes reniform,
of normal size and prominence. Antennae: Seg-
ment 1 clubshaped, bent, with 3 setae and fine
microsculpture of parallel lines; segment 2 about
as long as wide, segment 3 longer than wide, seg-
ments 4-7 transverse. Antennal clubs very large,
285
transverse reniform, entirely pubescent, their grea-
test (transverse) length 0.65 X the distance be-
tween the eyes on the vertex; no sutures visible
by pubescence, undersides with a large reniform,
flaccid impression; no sensorial plaques or pits
visible.
Pronotum: Two times wider than long in me-
dian line; base bisinuate, lateral sides moderately
convergent, near apex bent interiorly, anterolateral
angles blunt, emargination for the head moderately
deep. Marginal stria complete, though very fine
behind head. Foveolae behind eyes well-developed;
disk punctulate, along both sides a band of coarse
punctures, slightly impressed at apical onethird.
Coarse punctures along base; epipleura not fim-
briate. Scutellum triangular.
Elytra: Together 1.3 times wider than long, disk
and lateral sides punctulate, apical half moderately
punctate, punctation reaching humeral bosses in
interstria 1, and slightly entering interstriae 2-
4. Punctation towards apex smaller and denser,
reaching apical stria. Humeral bosses prominent.
Sides moderately convergent posteriorly, straight,
near the apex bent interiorly, apex of each elytron
roundedly truncate. Marginal epipleural stria dis-
tinct, ending shortly before posterolateral angle.
Marginal elytral stria distinct, continuous with
complete apical stria. Outer subhumeral stria short,
basal. Inner subhumeral stria medial, punctate,
reaching a little beyond dorsal stria 1, continuous
with distinct oblique humeral stria. Four dorsal
striae, punctate, ending a little beyond elytral mid-
286 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Figs. 1—5: Saprinus grandiclava nov.spec. 1: Body outline. 2: Pro- and mesosternum. 3: Aedeagus, right lateral view.
4: Aedeagus, dorsal view. 5: Eight sternite, ventral view. Scale lines: Left 1 mm. (figs. 1, 2), right 0.5 mm. (figs. 3—5).
dle, striae 1 and 3 a little longer than striae 2 and
4. Stria 4 connected in an arch with complete sutural
stria, these continuous with apical stria. Interstrial
distance 2 wider than interstrial distances 1 and 3.
Longitudinal strioles near base of stria 1 and 2.
Propygidium: Short, moderately punctate with
few punctules intermixed, finely microsculptured
between punctures; faint impressions laterally.
Pygidium: Punctured and microsculptured as
propygidium, punctures near apex smaller.
Prosternum (fig. 2): Internal prosternal striae
convergent between procoxae, next slightly diver-
gent anteriorly, then bent with blunt angles and
slightly convergent, at the apex united in a short
transverse arch. These striae gradually deeper an-
teriorly, sulciform, giving margins of prosternal
keel a convex ridgelike appearance. Prosternal keel
between striae slightly convex, minutely punculate.
No true lateral prosternal striae, these replaced by
descending ridges, which are deeply delimited be-
hind, and fuse with margins of prosternal keel at
position of the blunt angles. Antennal grooves
large and deep, encroaching upon prosternal keel,
prosteriorly delimited by the descending lateral
prosternal ridges, anteriorly not delimited by a
ridge. Grooves for the antennal funicles wide and
deep.
Mesosternum (fig. 2): Fine marginal stria com-
plete; anterior part finely punctate, posterior part
with coarse punctures, meso-metasternal suture
barely visible, straight, laterally obsolete.
Metasternum: Disk finely punctulate, before
posterior margin and along oblique lateral striae
with coarser punctures. Slightly impressed along
median line, especially anteriorly and posteriorly;
median line ending a little before hind margin.
Oblique lateral metasternal striae ending amply
before hind coxae, slightly sulciform anteriorly.
Abdomen: First visible abdominal sternite with
coarse punctures along anterior margin, smaller
punctures laterally and behind, disc finely punctu-
late. Oblique postcoxal striae ending a little before
hindmargin. Other sternites moderately punctate.
Legs: Protibiae with six teeth at outer margin, 4
additional minute denticles proximally, and 4 small
apical teeth. Meso- and metatibiae spinulate at su-
perior edge. Tarsae fimbriate inferiorly.
Genitalia: Aedeagus (figs. 3, 4): Slightly curved,
parameres not fused with penis, their tips round-
edly truncate. Eighth sternite: fig. 5.
KANAAR:
Holotype: “HOLLANDIA / NW. GUINEA / P. H. V.
DOESBURG”; "Museum Leiden / collectie / Van Does-
burg / rec. 1973”; red label: "Holotype @ / Saprinus /
grandiclava / P. Kanaar des. 1989”.
Holotype missing right mesotarsus, and 3 distal lids of
left mesotarsus and right metatarsus respectively. The
alae are stretched out. Allotype: Same data as holotype,
length: 4.3 mm. Width: 3.6 mm. Metasternum without
impression. Vaginal palpes deeply triangularly emargi-
nate at apex, styli long. Paratypes: 1 male, 2 females, same
data as holotype.
Variability: The divergency of the prosternal
striae varies a little; in one of the female paratypes
the striae are parallel, the blunt angles barely indi-
cated.
Holotype, allotype and one paratype in the col-
lection of the Rijksmuseum van Natuurlijke His-
torie in Leiden, two paratypes in the author's pri-
vate collection, which has been legated to the same
museum.
DISCUSSION
According to P. H. van Doesburg Jr. his father
has never been in Dutch New Guinea himself (per-
sonal communication). The beetles from Hollandia
and surroundings were collected on his request and
sent to him by one of his friends, Mr. W. Stüber.
Nothing is known about the date of the capture or
the circumstances under which the beetles were
found. Details concerning the collecting-activities
of Mr. Stüber are given by Lieftinck (1932).
The new species belongs to a group of Australian
and Pacific species, in which the paramera are not
fused with the penis. This is considered to be a
primitive feature (Dahlgren 1971). The absence of
sensorial plaques or pits on the antennal clubs
might be a primitive character too. Saprinus gran-
diclava differs among other things from all other
New Saprinus 287
members of this group by the different form of the
paramera and 8th sternites, which were pictured by
Dahlgren (1962, 1968, 1971). The antennal clubs
are unusually big; in the other members of this
group the clubs are of normal proportions in
Saprinus pseudocyaneus White, S. australis Boisdu-
val (syn.: S. tasmanicus Marseul) and S. detritus
Fabricius (syn. S. antipodus Dahlgren); in the de-
scriptions of the other species of the group, not
known to me (S. amethystinus Lewis and S. tyrrhe-
nus Blackburn), nothing is said about the dimen-
sion of the clubs, apparently being of normal pro-
portions.
Note: The name of the species is based on the
strikingly big antennal clubs.
ACKNOWLEDGEMENT
I thank Mr. J. Krikken for the opportunity to
study and describe the species, and for critically
reading the manuscript.
REFERENCES
Dahlgren, G., 1962. Über einige Saprinus-Arten (Col.
Histeridae). - Opuscula Entomologica 27: 237—248.
Dahlgren, G., 1968. Beiträge zur Kenntnis der Gattung
Saprinus (Col. Histeridae) III. - Entomologisk Tidsk-
rift 89: 255—268.
Dahlgren, G., 1971. Zur Taxonomie der Gattungen Sapri-
nus, Hypocaccus und Zorius (Col. Histeridae). - En-
tomologisk Tidskrift 92: 43—53.
Lieftinck, M. A., 1932. The Dragonflies (Odonata) of
New Guinea and neighbouring Islands. - Nova Gui-
nea 15: 485-602.
Received: 31 March 1989
Accepted: 17 July 1989
288 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
RECTIFICATION
The editors regret that the following table was accidentally omitted in the previous issue from: Th.
G. H. Platel, The egg laying and larval development of Pholcus phalangioides (Fuessling) (Araneae:
Pholcidae), Volume 132: 135-147.
Table 4. Characteristics of the first post-embryonic stages with Pholcus phalangiotdes. A comparison of
the terminology of Vachon (1957) and Canard (1984, personal statement 1985).
Postembryonic development
Embryonic
ÿ Larval period nympho-imaginal period
A | Development
5 prelarva (PL) prenymph (PN) first nymph (NI)
O
N hatching
moult
Shedding of
the egg-
Reversion Breaking of Breaking of membranes
the eggmem- che first
branes integument
= (chorion +
ayıng vitellina ;
membrane) Shedding of
the first
integument
hatching
moult 1
Pullus Projuvenile Juvenile
C Incomplete spiderling 1 (Jil) Complete spiderling (J2)
A
N Embryonic
È Juvenile period
development
R
D
Postembryonic development
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TIJDSCHRIFT VOOR ENTOMOLOGIE
DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING
UITGEGEVEN DOOR
MC7
IS GRAR Y
SEP 07
REGISTER VAN DEEL 132 / Index to volume 132
Een sterretje duidt een naam aan die nieuw is voor de wetenschap; cursieve paginanummers
verwijzen naar de beschrijvingen van nieuwe taxa.
An asterisk denotes a name new to science; numbers in italics refer to descriptions of new taxa.
ARANEAE
phalangioides, Pholcus 288
COLEOPTERA
amethystnus, Saprinus 287
antipodus, Saprinus 287
assimile, Agonum 244, 245
australis, Saprinus 287
biguttatus, Notiophilus 244, 245,
brevicollis, Nebria 244, 245
brunnea, Amara 244, 245, 248
Calyce 107
caraboides, Cychrus 244, 245
chinensis, Callosobruchus 154
coriaceus, Carabus 244, 245, 247,
[248
cristatus, Pterostichus 244, 245,
[248
detritus, Saprinus 287
gemellata, Mordellistena 105,
[106
granarius, Sitophilus 154
*grandiclava, Saprinus 285-287
Guineapolemius 129
*hammondi, Polemiosilis 130, 131
*leveyi, Mordellistena 105, 106
lindbergi, Mordellistena 106
madidus, Pterostichus 248
micropterus, Calathus 244, 245
Mordellistena 105
*nagaii, Polemiosilis 130, 133
*nanggalaensis, Polemiosilis 130,
[133
nemoralis, Carabus, 244, 245, 248
nitens, Trichotichnus 244, 245,
[248
oblongopunctatus, Pterostichus
1244, 245, 247
obscuricolor, Polemiosilis 131
parallelepipedus, Abax 244, 245,
1247, 248
parallelus, Abax 244, 245
placidus, Trichocellus 244,
Polemiosilis 129
problematicus, Carabus 244, 245,
[247, 248
pseudocyaneus, Saprinus 287
pumicatus, Stomis 244, 245
quadripunctatus, Harpalus 244,
[245, 248
rotundicollis, Calathus 244, 245,
[248
*rubidiceps, Polemiosilis 130,
[131-134
rufipes, Notiophilus 244, 245,
[248
rufomarginatus, Leistus 244, 245
rufosuta, Polemiosilis 129,
[131-133
rufosuta saleierensis,
| Polemiosilis 129, 132
saleierensis, Polemiosilis 129, 132
Saprinus 285
*simplex, Polemiosilis 129, 130
Sitophilus 154
spinibarbis, Leistus 244, 245, 248
*sumatrensis, Calyce 105, 106, 107
tasmanicus, Saprinus 287
testaceobasalis, Guineapolemius
[129, 134
testaceobasalis, Polemiosilis 129,
[134
tolianus, Guineapolemius 129,
[134
tyrrhenus, Saprinus 287
vernalis, Pterostichus 248
violaceus, Carabus 244, 245, 247,
[248
DIPTERA
Calliphora 154
domestica, Musca 154
Drosophila 152, 154
lucens, Lipara 154
melanogaster, Drosophila 152
Semudobia 154
Zaprionius 152
HETEROPTERA
abietis, Eremocoris 4, 28
abietis, Gastrodes 4
abietum, Gastrodes 4, 28
abutilon abutilon, Stictopleuris 32,
[81, 82
Acalypta 11
Acanthia 3
Acanthosoma 34
Acetropis 21
Acompocoris 23
Acompus 28
acuminata, Aelia 33
acuteangulatus, Gonocerus 30, 80,
[81
Adelphocoris 20
adenocarpi, Orthotylus 18
adspersus, Lygus 1, 2, 19, 56
Aelia 33
Aellopus 29
aeneus, Eusarcoris 5
aeneus, Eysarcoris 5, 33
aequale, Camptozygum 3, 20
aestivalis, Aphelocheirus 2, 9, 37,
[38
aestivalis, A phelochirus 2
Aethus 32
affinus, Calocaris 20, 61
affinis, Corixa 8
affinis, Scolopostethus 28
agilis, Dicranocephalus 30, 79
Agnocoris 20
Agramma 12
albicinctus, Psallus 15
albidus, Amblytylus 14
albipennis, Plagiognathus 15
alboacuminatus, Aphanus 4
alboacuminatus, Raglius 4, 29, 78
albolineatus, Cremnocephalus 16,
[53
albomaculatus, Melanocoryphus
[88
albomarginatus, Gnatoconus 5
alienus, Isometopus 65
290
Alloeotomus 13
alnicola, Psallus 3, 49
alpestris, Cremnocephalus 2
alpinus, Acompocoris 23, 69, 70
alpinus, Plagiognathus 15, 50, 51
Alydus 31
ambiguus, Psallus 15
Amblytylus 14
ambulans, Mecomma 19, 55, 56
ampliasta, Tingis 12
amplicollis, Anthocoris 23, 68, 88
Anaptus 22, 67
Aneurus 25
angulatus, Blepharidopterus 18,
[55
angusticollis, Peritrechus 29, 78
angustulus, Ischnocoris 28
annulatus, Dicyphus 17
annulatus, Harpactor 73
annulatus, Rhynocoris 4, 25, 71,
[73
annulicorne, Campylomma 3, 16,
51,32
annulicornis, Campylomma 3
annulipes, Deraeocoris 13, 44, 45
antennatus, Megalonotus 29, 78
antennatus, Rhyparochromus 78
antennatus, Teratocoris 22
Anthocoris 23
Aphanus 4, 30
Aphelocheirus 9
apterus, Halticus 17
apterus, Himacerus 4, 22
apterus, Nabis 4
apterus, Pyrrhocoris 30
Aptus 22, 67
Aquarius 2, 10
Aradus 25
arbustorum, Plagiognathus 15
Arctocorisa 2, 8
arenarius, Trapezonotus 30
arenicola, Saldula 11, 40
Arenocoris 31
argentatus, Gerris 10
Arma 34
Asciodema 16
asper, Gerris 2, 39
assimilis, Psallus 15, 49, 87
ater, Capsus 21
ater, Dufouriellus 4, 24
ater, Geocoris 27, 75, 87
ater, Xylocoris 4
aterrimus, Brachypelta 5
aterrimus, Cydnus 5, 32, 83
atomarius, Pinalitus 19, 57
Atractotomus 15
atrata, Microtoma 4
atratus, Aellopus 4, 29, 78
aurescens, Cymus 4, 27, 75
austriaca, Eurygaster 32, 83, 84
avenius, Aneurus 25, 74
baccarum, Dolycoris 33
baerensprungi, Empicoris 24, 72
basalis, Orthops 19, 57, 58
Bathysolen 31
beckeri, Megacoelum 20, 61
Beosus 29
Berytinus 4, 26
Berytus 4
betuleti, Psallus 15
biclavatus, Calocoris 20, 61
bicolor, Sehirus 32
bicuspis, Tetraphleps 23, 69
bidens, Picromerus 34
bidentata, Pygolampis 25, 73
biguttatus, Sehirus 32
bilineatum, Neomecomma 3, 18
bilineatus, Orthotylus 3
binotatus, Stenotus 21
bipunctata, Loricula 13
Blepharidopterus 18
bohemani, Microsynamma 3
bohemanni, Monosynamma 3, 16,
[52
bonsdorffi, Cymatia 2
bonsdorffii, Cymatia 2, 7
boops, Nabis 4
boops, Stalia 4, 22, 66, 67
Bothynotus 13
Brachyarthrum 14
Brachysteles 24
bevicollis, Amblytylus 1, 14, 47
brevipennis, Plinthisus 28
brevis, Nabis 23, 66, 67
brunneeus, Drymus 28
Bryocoris 13
buenoi, Microvelia 10, 38, 39
butleri, Anthocoris 23, 68
caelestialium, Trigonotylus 1, 2,
[22, 64, 65, 86
caerulea, Zicrona 34
c-album, Saldula 11, 40
calcarata, Stenodema 4, 21
calcaratum, Stenodema 4
calcaratus, Alydus 31
Callicorixa 8
Calocoris 20
campestris, Lyctocoris 4, 24
campestris, Orthops 19
Camptobrochis 3
Camptozygum 20
Campylomma 16
Campyloneura 17
Campylosteira 11
capitatum, Piesma 1
caprai, Velia 2, 10
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Capsodes 4, 21
Capsus 21, 62
capucina, Lasiacantha 12, 43
Cardiastethus 24
cardui, Tingis 12
caricis, Cyrtorhinus 3, 18, 56
caricis, Cyrtorrhinus 3
carinata, Acalypta 11
carinata, Acetropis 21
Carpocoris 33
castanea, Hesperocorixa 8
Catoplatus 12
cavifrons, Glaenocorisa 2
Ceraleptus 31
Ceratocombus 7
cervinus, Pinalitus 19, 57
Charagochilus 20
Chartoscirta 11
Chilacis 27
Chiloxanthus 10
chiragra, Megalonotus 29
Chlamydatus 15
chlorizans, Malacocoris 17
Chlorochroa 33, 85
Chorosoma 31
chrysanthemi, Plagiognathus 15
ciliata, Monanthia 43
Cimex 24
cimicoides, Ilyocoris 2, 8
cimicoides, Naucoris 2
cincta, Chartoscirta 11
cinerea, Nepa 7
cinnamomeus, Aradus 25
cinnamopterus, Pilophorus 17
clavatus, Pilophorus 17
claviculus, Cymus 27
clavipes, Berytinus 26, 88
coccinea, Pseudoloxops 3
coccineus, Pseudoloxops 3, 18
cocksii, Chartoscirta 11
coleoptrata, Cymatia 7
coleoptrata, Myrmedobia 13
coleoptratus, Ceratocombus 7
columbarius, Cimex 24, 72
Compsidolon 15
concinna, Callicorixa 2
concinna, Paracorixa 2, 8
concolor, Orthotylus 18
confusus, Anthocoris 23
confusus, Drymus 4, 76
confusus, Pilophorus 17
confusus, Psallus 3, 15, 48, 49
Conostethus 14
constrictus, Dicyphus 16, 53
contaminatus, Lygocoris 19
contractus, Taphropeltus 28, 77
convergens, Dictyla 3, 12
Coranus 25
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
cordiger, Deraeocoris 13
Coreus 5, 30
coriaceus, Orthocephalus 3, 17
Coriomeris 31
Corixa 8
Corizus 5, 31
corticalis, Aradus 25, 73
coryli, Phylus 14
costalis, Camptotelus 76
costalis, Tropidophlebia 4, 27, 76
costata, Physatocheila 12, 43
crassicornis, Criocoris 16, 52
crassicornis, Stictopleurus 1
crassipes, Berytinus 26
crassipes, Phymata 25, 73, 87
Cremnocephalus 16
Criocoris 16
crispata, Tingis 12, 43
cruciatus, Globiceps 3, 18
culiciformis, Empicoris 24
currens, Velia 2
cursitans, Sciocoris 33
cursitans, Xylocoris 24
custos, Arma 34
Cydnus 32
cylindricollis, Orthonotus 1
Cyllecoris 17
Cymatia 7
Cymus 27
Cyphostethus 34
Cyrtorhinus 18
Cyrtorrhinus 3
dahlmannii, Spathocera 5, 31
dalmani, Spathocera 5
decolor, Lopus 3, 14
decolor, Onychumenus 3
decoratus, Scolopostethus 28
denticulatus, Coriomeris 31
dentipes, Corixa 8
depressus, Aradus 25
depressus, Orsillus 26, 74, 88
Deraeocoris 3, 13
Derephysia 12
desertus, Trapezonotus 30, 79
diaphanus, Blepharidopterus 3,
[18,55
diaphanus, Orthotylus 3, 55
Dichrooscytus 20
Dicranocephalus 4
Dictyla 3, 12
Dictyonota 12
Dicyphus 16
dilatatus, Megalonotus 29
dimidiatus, Phytocoris 21
diminutus, Psallus 3, 49
dispar, Trapezonotus 30
dissimilis, Cimex 24, 72
distincta, Sigara 8
distingendus, Peritrechus 1
distinguenda, Myrmedobia 13, 45
ditomoides, Metopoplax 27, 75
dolobrata, Leptopterma 4, 22
dolobratus, Miris 4
Dolycoris 33
dominulus, Eurydema 34, 86
dorsalis, Odontoscelis 5, 83
Drymus 28
Dryophilocoris 18
dubius, Sehirus 1
Dufouriellus 24
dumetorum, Physatocheila 12, 44,
[45
dumosa, Jalla 34, 86
dumosus, Asopus 86
echii, Dictyla 12
Elasmostethus 34
Elasmucha 34
Elatophilus 23
elegantula, Chartoscirta 11, 41
elegantula, Loricula 13
elongata, Notostira 4, 21
Emblethis 30
Empicoris 4, 24
enervis, Lasiosomus 29, 77
Enoplops 30
epilobii, Dicyphus 16
equestris, Lygaeus 4, 26, 74
equestris, Spilostethus 4
Eremocoris 28
ericae, Ischnorhynchus 4
Ericae, Nysius 26
ericetorum, Nabis 23
ericetorum, Orthotylus 18
errans, Dicyphus 16
erratica, Notostira 4
erraticus, Eremocoris 4
erythrocephalus, Halticus 53
Eurydema 34
Eurygaster 32
evanescens, Chlamydatus 16, 50,
[51
exilis, Myrmedobia 3, 13
Eysarcoris 33
fabricii, Arctocorisa 2
fabricii, Catoplatus 12
fabricii, Eusarcoris 5, 33
falleni, Psallus 15
falleni, Pseudophloeus 5
falleni, Sigara 8
fallenii, Arenocoris 5, 31
fasciiventris, Cardiastethus 24, 71
favosus, Neides 1
ferrarii, Orthocephalus 1, 54
ferrugata, Elasmucha 34
ferrugata, Leptopterna 4, 22
ferrugatus, Miris 4
291
ferrugineus, Gastrodes 4
ferus, Nabis 23
fieberi, Elasmucha 34
Fieberocapsus 17
filicis, Monalocoris 13
flavellus, Psallus 15
flaveolus, Eurycolpus 1
flaveolus, Cyrtorrhinus 3
flaveolus, Fieberocapsus 3, 17, 54
flavicorne, Cydnus 5
flavicornis, Aethus 5, 32
flavilinea, Deraeocoris 1, 2, 13, 45,
[87,94
flavinervis, Orthotylus 18
flavomaculatus, Globiceps 18
flavomarginata, Nabicula 4
flavomarginatus, Nabicula 22
flavomarginatus, Nabis 4
flavoquadrimaculatus, Cyllocoris 3
flavoquadrimaculatus,
[Dryophilocoris 3, 18
flavosparsus, Orthotylus 18
foliacea, Derephysia 12
formicetorum, Xylocoris 24, 70,
[71
fossarum, Sigara 8
fracticollis, Pachybrachius 4, 29
fracticollis, Pamera 4
fucicola, Saldula 11, 40
fuligineus, Stygnocoris 29
fuliginosa, Dictyonota 12
fuliginosa, Odontoscelis 32
fulvicollis cruciatus, Globiceps 3,
[18
fulvipennis, Plagiognathus 15, 50,
[51
fulvomaculatus, Calocoris 20
furcata, Mesovelia 9
furcata, Notonecta 2
fuscescens, Orthotylus 18, 55, 88
fusciloris, Plagiognathus 3, 51
fuscispinus, Carpocoris 33, 86
galactinus, Xylocoris 24
Galeatus 12
galgulina, Phimodera 83
gallarumulmi, Anthocoris 4, 23
gallarum-ulmi, Anthocoris 4
Gampsocoris 26
Gastrodes 28
gemellatus, Lygus 19, 56, 57
geminata, Chartoscirta 1
geminus, Tytthus 1, 2, 14, 47, 87
geniculatus, Peritrechus 29
genistae, Heterocordylus 1, 2, 18,
[55
Geocoris 27
geoffroyi, Corixa 2
Geotomus 1
292
germanicus, Alloeotomus 13
germari, Arctocorisa 8
Gerris 2, 10
gibbifer, Gerris 10
gimmerthali, Acetropis 4
gimmerthalii, Acetropis 4, 21, 65,
[87
Glaenocorisa 7
glandicolor, Cymus 27
glauca, Notonecta 9
Globiceps 18
globulifer, Dicyphus 17
Gonianotus 30
Gonocerus 30
gothicus, Alloeotomus 13
gothicus, Capsodes 21, 62
gracilenta, Hydrometa 10
gracilis, Acalypta 11
gracilis, Myrmecoris 19, 55, 56
gracilis, Temnostethus 23
Graphosoma 33
Graptopeltus 29
grisea, Elasmucha 34
griseola, Micronecta 1
griseus, Conostethus 3, 14
griseus, Emblethis 30, 79
griseus, Pachymerus 79
grossipes, Gastrodes 4, 28
grylloides, Geocoris 27
guttula, Prostemma 22, 67
gyllenhali, Charagochilus 3
gyllenhalii, Charagochilus 3, 20
Hadrodemus 20
haematodes, Psallus 3, 15
haemorrhoidale, Acanthosoma 34
Hallodapus 16
Halosalda 11
Halticus 17
hamulatus, Taphropeltus 28, 77
Harpocera 14
Hebrus 9
hellensi, Arctocorisa 2
hellensii, Sigara 2, 8, 36
helveticus, Nysius 4, 26
Hesperocorixa 2, 8
Heterocordylus 18
Heterogaster 27
Heterotoma 18
hieroglyphica, Arctocorisa 2
Himacerus 22, 67
hirsutulus, Phytocoris 1
hirticornis, Berytinus 26
hirundinis, Oeciacus 24, 71, 72
histrionicus, Cyllecoris 3, 17
histrionicus, Cyllocoris 3
Holcostethus 5, 33
holesericeus, Polymerus 20, 59, 60
holosericeus, Tropistethus 28, 76,
[77
holsata, Stenodema 4, 21
holsatum, Stenodema 4
Hoplomachus 14
hottentotta, Eurygaster 3
hottentotta, Tetyra 83
humeralis, Phimodera 32, 83
humuli, Dictyla 3, 12
humuli, Monanthia 3
hyalinus, Liorhyssus 31, 81
Hydrometra 10
hyosciami, Therapha 5, 82
hyoscyami, Corizus 5, 31, 80, 81
Hypseloecus 17
iactans, Sigara 8, 36, 87
Ilyocoris 8
imitator, Micracanthia 41
infusum, Megacoelum 20
insignis, Phytocaris 21
intermedia, Rubiconia 33, 84
intermedius, Dichrooscytus 20, 60
interstinctus, Elasmostethus 34
intricatus, Phytocoris 21, 61, 62
intrusus, Isometopus 22, 65
inuncta, Podops 33
iracundus, Rhinocoris 4, 73
Ischnocoris 28
Ischnodemus 27
Isometopus 22
italicum, Graphosoma 5
Jalla 34
juniperi, Gonocerus 30, 80, 81
juniperi, Phytocoris 1
juniperina, Chlorochroa 33
Kalama 12
kalmi, Lygus 3
kalmii, Orthops 3, 19
Kleidocerys 27
klugii, Aelia 33
lacustris, Gerris 10
laeta, Serenthia 3
laetum, Agramma 3, 12
laevigata, Stenodema 4, 21
laevigatum, Stenodema 4
laevis, Aneurus 5, 25, 73, 74
Lamproplax 28
Lasiacantha 12
Lasiosomus 29
lateralis, Gerris 2, 10, 38, 39
lateralis, Haldosalda 3
lateralis, Halosalda 3, 11
lateralis, Sigara 2, 8
laticollis, Orius 23, 69, 70
lativentris, Nabis 4
latus, Drymus 4, 28, 76
lectularius, Cimex 24
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Legnotus 32
lepidus, Psallus 15
leptocerus, Heterocordylus 18, 54,
[55
Leptopterna 4, 22
leucocephalus, Halticus 54
leucocephalus, Strongylocoris 17,
[53, 54
limbata, Nabicula 4
limbatus, Anthocoris 23
limbatus, Lygocoris 19, 59
limbatus, Nabicula 22
limbatus, Nabis 4
limbosus, Legnotus 5, 32
limitata, Sigara 8
limitatum, Brachyarthrum 1, 2, 14
[49, 87
Limnoporus 10
linearis, Megaloceroea 4
linearis, Ranatra 7
lineata, Nabicula 4
lineatum, Graphosoma 5, 33, 84
lineatus, Nabicula 22
lineatus, Nabis 4
lineatus, Nysius 4
lineatus, Teratocoris 65
lineola, Odontoscelis 5, 32, 83
lineolatus, Adelphocoris 20
lineolatus, Camptotelus 4, 76
linnaei, Hesperocorixa 2, 8
linnei, Arctocorisa 2
Liocoris 19
Liorhyssus 31
litoralis, Plagiognathus 3, 15, 50,
[51
littoralis, Salda 11
lituratus, Piezodorus 33
lividus, Ceraleptus 31
longipalis, Sigara 8, 36, 37, 87
longirostris, Temnostethus 1, 2,
[23, 67
Lopus 4, 14
Loricula 3, 13
lucorum, Lygocoris 19
luctuosus, Sehirus 32
lugubris, Arctocorisa 2
lundii, Peritrechus 4, 29, 78
lurida, Pamera 4
luridus, Pachybrachius 4, 29, 78
luridus, Plociomerus 78
luridus, Psallus 15
luridus, Strongylocoris 17
luridus, Troilus 34
lutea, Notonecta 9, 37
luteicollis, Halticus 17, 53
lutescens, Deraeocoris 13
Lyctocoris 24
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Lygaeus 26
Lygocoris 3, 19
Lygus 3, 19, 57
lynceus, Aphanus 4
lynceus, Graptopeltus 4, 29
m-flavum, Hadrodemus 3, 20, 60,
[61
m-flavum, Homodemus 60
M-flavum, Homodemus 3
Macrodema 30
Macrolophus 16
Macrotylus 14
maculata, Notonecta 9
maculata, Piesma 4
maculatum, Piesma 4, 25
maculatus, Galeatus 12, 43, 87
maculatus, Naucoris 9, 37
maculatus, Rhopalus 31
maerkelii, Pithanus 19
magnicornis, Atractotomus 15, 51
major, Anaptus 4, 22
major, Nabis 4
majuscula, Triphleps 4
majusculus, Orius 4, 23
Malacocoris 17
mali, Atractotomus 15
marginalis, Micracanthia 11, 41,
[42
marginalis, Orthotylus 18
marginalis, Salda 41
marginata, Acalypta 11, 42
marginatus, Coreus 5, 30
marginatus, Syromastus 5
marginellus, Calocoris 60
marginepunctatus, Gonianotus 30
maritimum, Monosynamma 16
maritimus, Beosus 29
maritimus, Lygus 19, 57
marqueti, Reuteria 1, 2, 17, 54, 88
maura, Eurygaster 32
Mecomma 19
medius, Dicranocephalus 30, 79
megacephalus, Geocoris 27, 75
Megacoelum 20
Megaloceroea 21
Megalocoleus 14
Megalonotus 4, 29
melanocephalus, Cymus 27, 75
melanocephalus, Eusarcoris 5
melanocephalus, Phylus 14
Melanocoryphus 88
melanoscela, Salda 40
melanoscela, Saldula 11, 40
meridionalis, Micronecta 35
meriopterum, Heterotoma 3, 55
Mesovelia 9
Metatropis 26
Metopoplax 27
Micracanthia 11
Micronecta 7
Microphysa 3
microptera, Macrodema 30
Microsynamma 3
Microvelia 10
minki minki, Anthocoris 23, 68,
[69
minor, Berytinus 26, 74
minor, Plesiocoris 19, 58, 59
minuta, Triphleps 4
minutissima, Micronecta-7, 35, 36
minutissima, Plea 9
minutus, Orius 4, 23, 69
Miridius 21
miriformis, Myrmus 31
Miris 20
mirmicoides, Aptus 4, 22
modestus, Oxycarenus 27, 75, 76
moesta, Hesperocorixa 8
molliculus, Megalocoleus 14
mollis, Psallus 15, 49
Monalocoris 13
Monanthia 3 =
moncreaffi, Orthotylus 18
Monosynamma 3, 16
montivagus, Berytinus 26, 74
morio, Salda 11, 40
morio, Sehirus 32, 83
muelleri, Salda 11, 40
musci, Acalypta 11, 42
mutabilis, Orthocephalus 3
Myrmecoris 19
Myrmedobia 13
Myrmus 31
Nabicula 22, 67
Nabis 4, 23, 67
najas, Aquarius 10
nassatus, Orthotylus 18
nasutus, Amblytylus 14, 47
Naucoris 9
nebulosa, Rhaphigaster 1
nebulosus, Sphragisticus 29
Neides 26
nemoralis, Anthocoris 23
nemorum, Anthocoris 23
Neomecomma 18
Neottiglossa 33
Nepa 7
niger, Orius 4, 23
nigra, Triphleps 4
nigricornis, Carpocoris 86
nigricornis, Elatophilus 23, 67, 68,
[88
nigrina, Acalypta 12, 42, 43
nigrita, Aethus 32, 82, 83
nigrita, Cydnus 83
nigrita, Polymerus 20, 60
293
nigritulum, Monosynamma 1, 52
nigrolineata, Sigara 2, 8
nigrolineata, Trigonosoma 84
norvegicus, Calocoris 20
Notonecta 9
Notostira 21
nowickyi, Ptytocoris 62
nubilus, Bathysolen 31, 80, 81
nubilus, Macrolophus 16
nubilus, Peritrechus 29
Nysius 26
oberti, Stephanitis 12
obliqua, Notonecta 2, 9
obliquus, Cymus 4, 75
obscurellus, Phoenicocoris 3, 15
obscurellus, Psallus 3
obsoletum, Asciodema 16, 52
ochromelas, Calocoris 3
odontogaster, Gerris 10
Odontoscelis 32
Oeciacus 24
oleracea, Eurydema 34
olivaceus, Deraeocoris 13
Oncochila 12
Oncotylus 14
opacula, Saldula 11, 41
Orius 4, 23
ornatum, Eurydema 5, 86
Orsillus 26
Orthocephalus 17
orthochila, Saldula 11
Ortholomus 26
Orthonotus 14
Orthops 3, 19, 57
Orthotylus 18
Oxycarenus 27
pabulinus, Lygocoris 19
Pachybrachius 29
Pachycoleus 7
Pachytomella 17
palliceps, Phylus 14
pallicornis, Dicyphus 3, 16
pallidicornis, Dicyphus 3
pallidipennis, Ophthalmicus 75
pallidus, Dicyphus16
pallipes, Saldula 11, 41
Palomena 33
paludum, Aquarius 10
paludum, Teratocoris 22, 65
palustris, Polymerus 20
palustris, Saldula 1, 11, 41
Pantilius 21
panzeri, Corixa 8
Paracorixa 8
parallela, Pachytomella 17
parumpunctatus, Rhopalus 31
parvicornis, Brachysteles 24, 70,
(71
294
parvula, Acalypta 12
parvulus, Atractotomus 15, 49, 51
parvulus, Xylocoris 1
paykulit, Macrotylus 3
paykulli, Macrotylus 3, 14
pedestris, Stygnocoris 4
Pentatoma 33
Peribalis 5
Peritrechus 29
perplexus, Pilophorus 17
perrisi, Psallus 15
personatus, Reduvius 25
Phimodera 32
phoeniceus, Rhyparochromus 29
Phoenicocoris 15
Phylus 14
Phymata 25
Physatocheila 3, 12
Physatochila 3
Phytocoris 21
picea, Lamproplax 4, 28, 77
piceus, Drymus 4, 77
picinus, Gnatoconus 77
picipes, Legnotus 5, 32
Picromerus 34
pictus, Scolopostethus 28
Piesma 25
Piezodorus 33
Piezostethus 4
pilicornis, Calocoris 20, 61
pilicornis, Drymus 28, 76
pilifer, Capsus 1, 2, 21, 62, 63
Pilophoprus 17
pilosa, Tingis 12, 43
pilosella, Saldula 11
pilosus, Anthocoris 23, 68, 69
pilosus, Bothynotus 13, 45
pilosus, Chiloxanthus 10
pilosus, Megalocoleus 14
pilosus, Scolopostetus 28, 77
Pinalitus 3, 19, 57
pinastri, Camptozygum 3
pinetalla, Plesiodema 3, 14
pinetellum, Plesiodema 3
pini, Phytocoris 21, 62
pini, Rhyparochromus 29
pinicola, Chlorochroa 33, 85
Pionosomus 30
pipistrellae, Cimex 72
pipistrelli, Cimex 24, 72
Pitedia 85
Pithanus 19
Plagiognathus 15
planicornis, Heterotoma 3, 18, 55
platycheila, Acalypta 3, 12, 42, 43
platychila, Acalypta 3, 42
Plea 9
plebejus, Eremocoris 28
Plesiocoris 19
Plesiodema 14
Plinthisus 28
Ploiariola 4
podagricus, Eromocoris 28, 77
Podops 33
Poeciloscytus 3
Polymerus 3, 20
populi, Lygocoris 1, 2, 19, 59
populi, Phytocoris 21
poweri, Micronecta 7, 35, 36
praetextatus, Megalonotus 29
praeusta, Callicorixa 1, 8
prasina, Palomena 33
prasinus, Orthotylus 18
pratensis, Lygus 19, 57
privignis, Kleidocerys 1, 27, 88
privignus, Kleidocerys 2, 74, 75
producta, Callicorixa 1
propinqua, Glaenocorisa 2, 7
Prostemma 22
Psallus 15
psammaecolor, Trigonotylus 22
pselaphiformis, Loricula 13
pseudoferus, Nabis 23, 66, 67
pseudograndis, Scolopostethus 1,
[2, 28, 77
Pseudoloxops 18
pseudoplatani, Psallus 15, 48, 49,
[87
pteridis, Bryocoris 13
Pterotmetus 30
puberulus, Scolopostethus 28
pulchella, Scolopocelis 1, 2, 24, 71,
[88
pulchellus, Trigonotylus 22
pulicarius, Chlamydatus 15
pullus, Chlamydatus 16
pumilio, Drymus 28, 76, 83
punctata, Corixa 2, 8
punctatonervosus, Stictopleurus
[32, 82
punctatus, Rhacognathus 34
punctipennis, Ortholomus 4, 26,
[74
punctipennis, Nysius 4, 74
punctipes, Gampsocoris 4, 26
punctipes, Metacanthus 4
punctipes, Oncotylus 14, 45, 46
punctulatus, Deraeocoris 13, 44,
[45
punctulatus, Geotomus 1
purpureipennis, Carpocoris 33
pusilla, Neottiglossa 33
pusillus, Hebrus 9
pusillus, Plinthisus 28, 76
pusillus, Psilophorus 88
pusillus, Temnostethus 23
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
pygmaea, Microvelia 1, 10, 39
pygmaeus, Acompocoris 23
pygmaeus, Cyrtorrhinus 47
pygmaeus, Tytthus 14, 47, 48
Pygolampis 25
pyrioides, Stephanitis 1
Pyrrhocoris 30
quadrata, Piesma 4
quadratum, Piesma 4, 25, 73, 74
quadratus, Aphanus 4
quadratus, Xanthochilus 4, 29
quadrimaculata, Psysatocheila 3,
143, 45
quadripunctatus, Adelphocoris 20
quadripunctatus, Calocoris 61
quadrivirgatus, Miridius 21, 61
quercus, Psallus 15
Raglius 29
Ranatra 7
reclairei, Agnocoris 3, 20, 58, 59
recticornis, Megaloceraea 4, 21,
[64, 65
Reduvius 25
resedae, Ichnorhynchus 4
resedae, Kleidocerys 4, 27
reticulata, Microvelia 2, 10
reticulata, Tingis 12, 43
reuteri, Notonecta 9, 37, 38
reuteri, Phytocoris 21
Reuteria 17
Rhacognathus 34
rhamnicola, Lygocoris 19
rhododendri, Stephanitis 12
rhombea, Verlusia 5
rhombeus, Syromastus 5, 30
Rhopalus 5, 31
Rhynocoris 25
Rhyparochromus 4, 29, 78
rivulorum, Velia 1, 2
rolandri, Aphanus 4, 30
rolandri, Calyptonotus 4
roseomaculatus, Calocoris 20
roseri, Salicarus 3, 16
roseri, Sthenarus 3
roseus, Conostethus 14, 46
roseus, Psallus 3
rotermundi, Sthenarus 3, 16
rottermundi, Sthenarus 3
ruber, Deraeocoris 13
Rubiconia 33
rubicundis, Agnocoris 20, 58, 59
rubicundus, Lygus 3
rubidus, Orthotylus 18
rubricatus, Pinalitus 19, 57
rufescens, Allodapus 3
rufescens, Hallodapus 3, 16, 53
rufescens, Metatropis 26, 74
rufescens, Pachycoleus 2, 35
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
ruficeps, Hebrus 9
ruficornis, Trigonotylus 22, 64, 65,
[86
rufifrons, Byrsoptera 3
rufifrons, Orthonotus 3, 14
rufipennis, Dichrooscytus 20
rufipes, Acompus 28
rufipes, Pentatoma 33
rufoscutellatus, Gerris 3
rufoscutellatus, Limnoporus 3, 10
rufus, Rhopalus 1
rugicollis, Plesiocoris 19
rugosus, Nabis 23
rugulipennis, Lygus 19
rusticus, Stygnocoris 29
ryeii, Drymus 1, 28, 77
sabuleti, Ischnodemus 27
sabulicola, Megalonotus 1, 29, 79
sabulicola, Monosynamma 16, 51,
[52
sabulosus, Stygnocoris 4, 29
sahlbergi, Hesperocorixa 8
Salda 11
Saldula 3, 11
Salicarus 16
salicellus, Compsidolon 3, 15
salicellus, Psallus 3
salicis, Psallus 3, 15, 49
salinus, Conostethus 3
salsolae, Piesma 25, 74
saltator, Halticus 17, 53
saltator, Orthocephalus 17, 54, 55
saltatoria, Saldula 11, 41
saltitans, Chlamydatus 16
sanguinipes, Pinthaeus 1
sarothamni, Anthocoris 23
saulii, Velia 10, 37
saundersi, Teratocoris 22, 64, 65
saxatilis, Lygnaeus 74
saxatilis, Spilostethus 26, 74, 87
scapha, Coreus 5
scapha, Enoplos 5, 30
scarabaeoides, Thyreocoris 32
schillingi, Chorosoma 31
schmidtii, Calocoris 20, 61
schneideri, Microvelia 2
scholtzi, Micronecta 7, 35
Sciocoris 33
Scoloposcelis 24
Scolopostethus 28, 77
scotica, Saldula 11, 39, 40
scotti, Sigara 8
scutellaris, Capsus 46
scutellaris, Deraeocoris 13, 45, 46
Sehirus 32
selecta, Sigara 8, 36
semistriata, Sigara 8
senecionis, Nysius 26
seticornis, Adelphocoris 20
sibiricus, Anthocoris 68
Sigara 2,8
signaticornis, Aradus 25, 73, 88
signoreti, Berytinus 26
simplex, Oncochila 12, 45
simulans, Anthocoris 23, 68, 69
singeri, Phytocoris 21, 62
smreczynskii, Physatocheila 3, 12,
[44, 45
solitarius, Macrotylus 14, 47
Spathocera 31
sphacelatus, Holcostethus 33, 84
sphacelatus, Holostethus 84
sphegiformis, Globiceps 18, 54
Sphragisticus 29
Spilostethus 4, 26
spinolae, Lygocoris 19
stagnalis, Sigara 2, 8
stagnorum, Hydrometra 10
Stalia 22, 67
staphiliniformis, Pterotmetus 4,
[30
staphylinoides, Pterotmetus 4 _
Stenocephalus 4
Stenodema 21
Stenotus 21
Stephanitis 12
Sthenarus 16
Stictopleurus 32
striata, Pycnopterna 3
striata, Sigara 8
striatellus, Calocoris 3, 20, 61
striatus, Miris 3, 20
strichnocera, Dictyonota 12
Strongylocoris 17
Stygnocoris 29
subapterus, Coranus 25, 72, 73
subpatellatus, Lopus 46
subrufus, Corizus 82
subrufus, Rhopalus 31, 81, 82
sulcatus, Capsodes 1, 2, 21, 62, 88
sulcicornis, Criocoris 1, 2, 16, 52,
[88
sylvaticus, Drymus 28
sylvestris, Peritrechus 4, 78
symphyti, Monanthia 3
Syromastus 30
Systellonotus 16
Taphropeltus 28
Temnostethus 23
tenella, Myrmedobia 3
tenellus, Orthotylus 18
Teratocoris 22
testudinaria, Eurygaster 32
Tetraphleps 23
thomsoni, Scolopostethus 28
thoracica, Harpocera 14
295
thoracicus, Gerris 10
thunbergi, Hoplomachus 14, 47
thunbergii, Lygus 47
thymi, Nysius 26
Thyreocoris 32
tibialis, Heterocordylus 18
ticinencis, Adelphocoris 20
tiliae, Phytocoris 21
Tingis 12
tipularius, Neides 26
Trapezonotus 30
tricornis, Dictyonota 3
tricornis, Kalama 3, 12
trifasciatus, Deraeocoris 13
Trigonotylus 22
triguttatus, Systellonotus 16
tripustulatus, Liocoris 19
trispinosa, Stenodema 4, 21, 63, 65
trispinosum, Stenodema 4
tristriatus, Cyphostethus 34
Troilus 34
Tropidophlebia 27
Tropistethus 28
truncatulus ericae, Kleidocerys 1,
[2, 4, 27,75, 88
tuberculifer, Coranus 1
tumidicornis, Heterocordylus 18,
[55
tunicatus, Pantilius 21
typhae, Chilacis 27
Tytthus 3, 14
ulmi, Phytocoris 21
umbricola, Microvelia 39
umbrinus, Sciocorus 1
unifasciatus, Polymerus 20
urticae, Heterogaster 27
vagabundus, Empicoris 24
variabilis, Psallus 15
varians, Psallus 15
varipes, Phytocoris 21
varius, Pionosomus 30
Velia 10
venator, Gonocerus 81
ventralis, Eurydema 1, 5, 34, 86, 87
venusta, Sigara l
venustus, Conostethus 14, 46, 47
verbasci, Campylomma 16
verbasci, Emblethis 30, 79
verna, Campylosteira 3, 11
verna, Campylostira 3
vernalis, Holcostethus 33, 84, 85
vernalis, Peribalus 84
vestita, Saldula 40
vicinus, Orius 23, 69
virens, Orthotylus 18, 56
virens, Stenodema 21, 63, 65
virescens, Orthotylus 18
virgula, Campyloneura 17
296
viridiflavus, Oncotylus 14, 46
viridinervis, Orthotylus 18
viridis, Lygocoris 19
viridis, Notonecta 9
viridissima, Palomena 33, 85
visci, Anthocoris 23, 69
visci, Hypseloecus 17, 53
viscicola, Pinalitus 19, 57
vitellinus, Plagiognathus 15
vulneratus, Polymerus 20, 59
wagneri, Capsus 21, 62, 63
wagneri, Lygus 1, 2, 19, 57
wagneri, Psallus 15, 48, 49
waltli, Pachycoleus 2, 7, 35
waltli, Pseudophloeus 5, 81
waltlii, Arenocoris 5, 31, 81, 87
waltlii, Pseudophloeus 81
wollastoni, Callicorixa 1
woodroffei, Coranus 1, 2, 25, 72,
[73
Xanthochilus 2
Xylocoris 4, 24
Zicrona 34
9
HOMOPTERA
Brachylobopyga 123, 127
citri, Planococcus 152
decorata, Brachylobopyga 123,
[124
Dilobopyga 123, 127
margarethae, Dilobopyga 127
minahassae, Dilobopyga 127
*montana, Brachylobopyga 123,
[124-127
toradja, Brachylobopyga 123-125,
[127
toradja, Cicada 123, 124
toradja, Cosmopsaltria 124
toradja, Tibicen 124
toradjus, Tibicen 124
HYMENOPTERA
Ablaxia 150, 151
aceris, Pediaspis 154
achaeus, Psilonotus 154
adamas, Psilonotus 154
Aggelma 150
Anisopteromalus 151
Anogmus 150, 151
Apantheles 154
basalis, Dinarmus 154
bedeguaris, Habrocytus 154
boarmiae, Dibrachys 154
Caenasis 150, 151
calandrae, Anisopteromalus 154
cavus, Dibrachys 154
Cecidostiba 150, 151
Cryptus 251, 252, 255, 257
dactylopii, Leptomastix 152
dianae dianae, Cryptus 251-253
dianae obscuripes, Cryptus 253
*dianae solivagus, Cryptus 251,
[252, 253
Dibrachys 151
Dinarmus 151
discus, Peridesmia 154
distinguendus, Lariophagus 154
divisa, Cynips 154
dubius, Mesopolobus 154
elevatus, Habrocytus 154
Eupteromalus 150
fasciiventris, Mesopolobus 154
fidenas, Vrestovia 152
*furvus, Cryptus 251, 252, 257
graminicola, Meraporus 154
Habrocytus 151
Hobbya 150, 151
hohenheimensis, Anogmus 154
Hypopteromalus 151
*insularis, Cryptus 251, 252, 256,
[258
Itamoplex 251
Lariophagus 151
lauta, Caenasis 154
liparae, Stenomalina 154
mediterraneus, Mesopolobus 154
Meraporus 151
Mesopolobus 150, 151
micropterus, Trichomalopsis
| (Eupteromalus) 154
muscarum, Pachyneuron 152, 153
Muscidifurax 151
Nasonia 150, 151, 154
nigricola, Toxeumorpha 152
nigritarsis, Cryptus 251, 252,
[254-256
obscuripes, Cryptus 251, 253, 254,
[257
*obscuripes impeditus, Cryptus
251,252, 21. 255
Pachycrepoideus 152
Pachyneuron 152, 153
Peridesmia 151
piceae, Anogmus 154
planiscuta, Pachyneuron 152
*praefortis, Cryptus 251, 252, 255
Psilonotus 150, 151
Pteromalus (Habrocytus) 151,
[153
punctipleura, Trychnosoma 154
puparum, Preromalus 153, 154
raptor, Muscidifurax 154
raptorellus, Muscidifurax 154
rhabdophagae, Mesopolobus 154
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
rotundata, Megachile 154
semifascia, Cecidostiba 154
sequester, Habrocytus 154
spinosus, Cryptus 255
Stenomalina 151
stenonota, Hobbya 154
strobilorum, Anogmus 154
subspinosus, Cryptus 255
Synedrus 151
tabacum, Hypopteromalus 154
tigasis, Trichomalopsis
[(Eupteromalus) 154
Toxeumorpha 152
Trichomalopsis 150, 151, 154
Trichomalus 151
Trychnosoma 151
*ultramondanus, Cryptus 251, 252,
[256, 257
uniraptor, Muscidifurax 154
vala, Anogmus 154
venustus, Pteromalus 154
verutus, Cryptus 251, 252, 257
verutus hierroensis, Cryptus 251,
[257
vindemmiae, Pachycrepoideus
[SI 252
vitripennis, Nasonia 151, 154
Vrestovia 151, 152
zaraptor, Muscidifurax 154
LEPIDOPTERA
adspersella, Coleophora 211
aequigesa, Coleophora 208
*alecturella, Coleophora 199, 200,
[204, 206, 217, 219, 231
algeriensis, Coleophora 201
artemisiella, Coleophora 200, 210
astragalella, Coleophora 203, 224
bagorella, Coleophora 200, 208
*batangica, Coleophora 199, 200,
[211, 217, 219, 240
*buteella, Coleophora 199-201,
(AOS NEN
*caradjai, Coleophora 199, 200,
[202, 203, 216, 218, 222, 223
canariipennella, Coleophora 204
changaica, Coleophora 200, 203
citrarga, Coleophora 200, 205, 216
Coleophora 200, 201
*cristata, Coleophora 199, 200,
[205, 206, 215, 218, 229
*denticulata, Coleophora 199, 200,
[210, 217, 219, 238
discordella, Coleophora 202
ditella, Coleophora 204
eothina, Coleophora 201
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
eteropennella, Coleophora 201,
[220
eurasiatica, Coleophora 200, 201
*falcipenella, Coleophora 199, 200,
1208; 215, 219; 235
galbulipennella, Coleophora 210
glaucicolella, Coleophora 205
*hoeneella, Coleophora 199, 200,
[203, 204, 216, 218, 225
hsiaolingensis, Coleophora 199,
[200, 207, 215
hungariae, Coleophora 210
immortalis, Coleophora 199, 200,
[205
issikii, Coleophora 210
kurokoi, Coleophora 200, 208, 216
laricella, Coleophora 204
*lucida, Coleophora 199, 200, 201,
[215, 218, 220
lunensis, Coleophora 207
mandschuriae, Coleophora 199,
[200, 203
melanograpta, Coleophora 199,
[200, 202, 218
microtitae, Coleophora 210
monoceros, Coleophora 207
neviusella, Coleophora 199
neviusiella, Coleophora 200, 201
nivifera, Coleophora 199, 200, 207
otitae, Coleophora 210
pandionella, Coleophora 210
parenthella, Coleophora 208
Pieris 154
*plurispinella, Coleophora 199,
(203, 215, 218, 224
pseudociconiella, Coleophora 210
pseudolinosyris, Coleophora 207
scioleuca, Coleophora 199, 200,
[206, 215, 219, 230
seminalis, Coleophora 199, 200,
(2055.27; 2185.2275.228
sinensis, Coleophora 199, 200, 204
*sittella, Coleophora 199, 200,
BO7 ZA GANS), ay
sternipennella, Coleophora 207
stramentella, Coleophora 203
summivola, Coleophora 199, 200,
[208, 217, 219, 233, 234
therinella, Coleophora 206
*tibetana, Coleophora 199, 200,
(204, 216, 218, 226
treskaensis, Coleophora 210
*tuberculata, Coleophora 199, 200,
(22108206 217; 219% 239
unipunctella, Coleophora 211, 212
versurella, Coleophora 199, 200,
[207
vestianella, Coleophora 200, 207,
[208
vestianella, Phalaena (Tinea) 207
vibicigerella, Coleophora 199,
[200, 203
virgaureae, Coleophora 206
weymarni, Coleophora 200, 211
yomogiella, Coleophora 200, 208,
[216
*yunnanica, Coleophora 199, 200,
120972177, 2192367237.
zelleriella, Coleophora 202
ODONATA
aglaia, Nannophlebia 121
apicalis, Palaemnema 167
appendiculatus,
[Phyllogomphoides 169
armatus, Epigomphus 158-160,
BSD
asiatica, Lathrecista 119
assimilis, Hemicordulia 118
*baltodanoi, Palaemnema 163-165,
[167
basiguttata, Gynacantha 118
batesi longipolex, Neocordulia
[163
bayadera, Gynacantha 118
berenice, Philogenia 168
bifasciatus, Phyllogomphoides
[169
*burgosi, Phyllogomphoides 163,
[168, 169, 171
capitatus, Paragomphus 121
celebense, Pseudagrion 117
Celebophlebia 115
Ceriagrion sp. nov. 1 & 2 117
chirripa, Cora 165
chrysis, Orthetrum 119
cinctus, Celebargiolestes 120
clavatus, Epigomphus 173
cleis, Lyriothemis 119
crepidus, Epigomphus 173, 174
congener, Potamarcha 119
*corniculatus, Epigomphus 158,
[159
cornutifrons, Phyllogomphoides
[169
crocops, Pseudagrion 117
delecolli, Celebothemis 120
Diplacina sp. nov. 1 & 2 119
distadens, Palaemnema 167
Drepanosticta sp. | & 2 118
*echeverrii, Epigomphus 163, 170,
[171-174
Epigomphus 165, 173, 174
297
eurybia, Tramea 120
femina femina, Agriocnemis 117
festiva, Trithemis 119
filostyla, Heliaeschna 118
flavescens, Pantala 120
flavipennis, Nososticta 118
fluctuans, Neurothemis 129
frontalis, Rhinocypha 117
glaucum, Orthetrum 119
helena, Philogenia 168
helvola, Teinobasis 117
*houghtoni, Epigomphus 163, 174,
[175
*indicatrix, Phyllogomphoides 155
[156
insignis quatuornotata,
[Agrionoptera 119
irina, Macromia 118
irregularis leptoptera,
[Tetrathemis 118
jaspidea, Anaciaeschna 118
majuscula, Hetaerina 165
militaris, Diplacina 119
monochroa, Rhinocypha 117
mysis, Nesoxenia 119
Neurothemis sp. nov. 119
Nososticta sp. nov. 1 & 2 118
kaupi kaupi, Neurobasis 116
lineata, Cratilla 118
obtusum, Zyxomma 119
occipitalis, Epigomphus 175
Ophiogomphus 173
pacificus, Phyllogomphoides 155
Palaemnema 165
pallidistylus, Perigomphus 163
paulirica, Palaemnema 167
paulsoni, Epigomphus 173
*peacocki, Philogenia 163,
[165-167, 168
pechumani, Epigomphus 173, 174
penelope, Gynacantha 118
Philogenia 168
Phyllogomphoides 155, 169
praemorsus, Lestes (Paralestes)
[117
Protosticta sp. 118
pruinosum, Orthetrum 121
pugnifer, Phyllogomphoides 169,
[171
pygmaea, Nannophya 118
rapax, Brechmorhoga 165
rogersi, Argia 163
rosenbergi, Gynacantha 118
rubescens, Argiocnemis 117
rufescens, Libellago 117
rufescens rufescens, Libellago 117
sabina, Orthetrum 119
298
sanguinolenta, Diplacina 119
Selysioneura 118
semicircularis, Phyllogomphoides
[155
senegalensis, Ischnura 117
servilia, Crocothemis 119
stigmatizans manadensis,
[Neurothemis 119
suasillus, Phyllogomphoides 155
subobtusus, Epigomphus 173, 175
subsimilis, Epigomphus 173, 175
superba, Teinobasis 117
tillarga, Tholymis 119
tincta semitincta, Rhinocypha 117
torrenticola, Diplacina 121
transmarina, Tramea 120
tristani, Erpetogomphus 163
trivialis, Diplacodes 119
tumefactus, Epigomphus 173, 175
ustum, Pseudagrion 117
xanthocyana, Libellago 177
ORTHOPTERA
*aulaeiadventa, Samariella 110, 111
bakeri, Samariella 110
celebesica, Karnydia 109, 110-113
cruciger, Ephippiger 177-180
cunii, Ephippiger 177-180
cunii f. jugicola, Ephippiger 178,
[180
dentata, Homeomastax 112
Ephippiger 177-180, 184, 185, 188
ephippiger, Ephippiger 177, 183
ephippiger diurnus, Ephippiger
[178-180
ephippiger vitium, Ephippiger
[177
ephippiger vicheti, Ephippiger
[177-180, 183, 184, 186
gracilipes, Karnydia 109, 112, 113
gracilis, Karnydia 110
Karnydia 109, 110, 113
*laticlava, Karnydia 110-113
milleri, Pseudommnesicles 109
*monki, Karnydia 110, 111, 113
plagiator, Pseudomnesicles 109
provincialis, Ephippiger 177-180
Pseudomnesicles 109
*rhodopeplus, Pseudomnesicles
[109-111
roseosignatus, Pseudomnesicles
[109, 110
rugosicollis, Uromenus 177-180
Samariella 109
terrestris, Ephippiger 177, 179,
[183, 184, 186, 187, 190-198
terrestris bormansi, Ephippiger
[177, 178, 180, 183-187, 190,
[191, 194-198
terrestris caprai, Ephippiger
[177-180, 183, 184, 186, 187, 190,
[191, 194-198
terrestris terrestris, Ephippiger
[177-180, 183, 184, 186, 187, 191,
[194, 196, 198
t. terrestris f. minor, Ephippiger
[178, 180, 187, 190, 194-197
Uromenus 178-180
viduata, Samariella 110
PLANTAE
Acer 154
Acer campestre 45
Acer pseudoplatanus 45
Achillae 203
Alnus glutinosa 75
Amaranthus 205, 207
Amaranthus paniculatus 205
Arctia 154
Artemisia 203, 210
Artemisia campestris 57
Artemisia princeps 208
TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 132, 1989
Artemisia montana 208
Artemisia vulgaris 57
Asparagus 154
Athyrium 112
Atriplex 207, 208
Betula 154
Buxus 68
Calluna vulgaris 75
Centaurea 154
Chamaecyparis lawsoniana 74
Chenopodium 207, 208
Chrysanthemum morifolium var.
[sinence 208
Cyathea 112
Cyathea multiflora 112
Dennstaedtia 112
Dennstaedtia obtusifolia 112
Diplazium 112
Diplazium expansum 112
Diplazium sp. aff. herbaceum 112
Elatostema 112
Fraxinus exselsior 68
Gnetum 109
Gnetum rhodopeplus 110
Larix principis-rupprechtii 205
Linaria vulogaris 154
Malus 201
Molinia caerulea 62
Nephrolepis 112
Phragmites 154
Picea abies 154
Populus 68
Populus canescens 59
Populus tremula 49
Quercus 154, 202
Rosa 154
Rubus 201
Salix 68
Salix repens 154
Thuja occidentalis 74
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