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Volume 133 1990
Tijdschrift
voor
Entomologie
A journal of systematic and evolutionary
entomology since 1858
oeloes
SNS
Se 0
m 2 ] 4
a HC i \
"> by
ON VE
RE 270)
CERN où
Published by the Nederlandse Entomologische Vereniging
Tijdschrift voor Entomologie
A journal of systematic and evolutionary entomology since 1858
Scope
The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a
long tradition in the publication of original papers on insect taxonomy and
systematics. The editors particularly invite papers on the insect fauna of the
Palaearctic and Indo-Australian regions, especially those including evolutionary
aspects e.g. phylogeny and biogeography, or ethology and ecology as far as
meaningful for insect taxonomy. Authors wishing to submit papers on disciplines
related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology
and applied entomology, are requested to contact the editorial board before
submitting. Usually, such papers will only be published when space allows.
Editors
E. |. van Nieukerken (elected 1986) and J. van Tol (1985)
Co-editors
A. W. M. Mol (1990) and R. T. A. Schouten (1990)
Advisory board
M. Brancucci (Basel), Y. Hirashima (Fukuoka) and M. R. Wilson (London).
The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the
‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society),
Amsterdam.
Editorial address
c/o National Museum of Natural History,
Postbus 9517, 2300 RA Leiden, The Netherlands.
Correspondence regarding membership of the society, subscriptions and
possibilities for exchange of this journal should be addressed to:
Nederlandse Entomologische Vereniging
c/o Instituut voor Taxonomische Zoölogie
Plantage Middenlaan 64
1018 DH Amsterdam
The Netherlands
Subscription price per volume Hfl. 300.- (postage included).
Special rate for members of the society. Please enquire.
Instructions to authors
May be obtained from the editors.
Graphic design
Ontwerpers B.V., Aad Derwort, ‘s-Gravenhage
Tijdschrift voor Entomologie
Contents of Volume 133
Articles
3
121
27
143
149
197
31
205
239
39
43
245
Asquith, A. & J. D. Lattin
Nabicula (Limnonabis) propinqua (Reuter) (Heteroptera: Nabidae): dimorphism,
phylogenetic relationships and biogeography.
Aukema, B.
Taxonomy, life history and distribution of three closely related species of the
genus Calathus (Coleoptera: Carabidae).
Bedos, A. & L. Deharveng
New species of Superodontella Stach (Collembola: Odontellidae) from Thailand.
Belle, J.
Progomphus nigellus and Phyllocycla hamata, two new dragonflies from Brazil
(Odonata: Gomphidae).
Belle, J. & J. van Tol
Anomalagrion hastatum (Say), an American damselfly indigenous to the Azores
(Odonata, Coenagrionidae).
Chaudhuri, P. K. & S. Chattopadhyay
Chironomids of the rice paddy areas of West Bengal, India (Diptera:
Chironomidae).
Mastrigt, H. J. G. van
New (sub)species of Delias from the central mountain range of Irian Jaya
(Lepidoptera: Pieridae).
Monk, K. A. & R. K. Butlin
A biogeographic account of the grasshoppers (Orthoptera: Acridoidea) of
Sulawesi, Indonesia.
Nieukerken, E. J. van
The Trifurcula subnitidella group (Lepidoptera: Nepticulidae): taxonomy,
distribution and biology.
Nieukerken, E. J. van
Stigmella rolandi sp. n.: a widespread southern European species on Rosa
(Lepidoptera: Nepticulidae).
Pape, T.
Two new species of Sarcophaga Meigen from Madeira and mainland Portugal
(Diptera: Sarcophagidae).
Pape, T.
Revisionary notes on American Sarcophaginae (Diptera: Sarcophagidae).
Pérez, T. M. & W. T. Atyeo
New taxa of feather mites (Acarina, Pterolichidae) from Megapodes (Aves,
Megapodiidae).
Quicke, D. L. J. & C. van Achterberg
The type specimens of Enderlein’s Braconinae (Hymenoptera: Braconidae)
housed in Warsaw.
265
75
269
85
97
273
107
Schouten, R. T. A.
A new species of Euchromius Guenée, 1845 (Lepidoptera: Pyralidae: Crambinae)
from Australia.
Sruoga, V.
Seven new species of Elachistidae (Lepidoptera) from the USSR.
Stock, J. H.
A new species of Austrodecus (Pycnogonida) from New South Wales, Australia.
Theowald, Br. & P. Oosterbroek
Zur Zoogeographie der Westpaläarktischen Tipuliden. IX. Die Tipuliden des
Vorderen Orients.
Tol, J. van
Key to the Malesian species of Leptogomphus Selys, with the description of a new
species from Sabah (Odonata, Gomphidae). Descriptions and records of Malesian
Odonata, |.
Tol, J. van
Zoological expeditions to the Krakatau Islands, 1984 and 1985: Odonata. Des-
criptions and records of Malesian Odonata, 2.
Wittmer, W.
Die Familie Malachiidae (Coleoptera) auf Sulawesi. 43. Beitrag zur Kenntnis der
indo-malaiischen Fauna.
Book announcements and reviews
96
268
268
268
268
R. Johansson et al., The Nepticulidae and Opostegidae (Lepidoptera) of North
West Europe. [R. T. A. Schouten]
T. B. Larsen, The butterflies of Egypt. [R. T. A. Schouten]
O. |. Merzheevskaya, Larvae of owlet moths (Noctuidae).
P. W. Oman, W. J. Knight & M. W. Nielson, Leafhoppers (Cicadellidae): a bibli-
ography, generic check-list and index to the world literature 1956-1985.
D. J. Williams & G. W. Watson, The scale insects of the Tropical South Pacific
region, 3.
Referees for volume 133
B. Aukema (Renkum), M. F. Claridge (Cardiff), R. W. R. J. Dekker (Amsterdam), Th. S.
van Dijk (Wijster), W. N. Ellis (Amsterdam), H.-J. Geiger (Bern), Th. Heijerman (Wage-
ningen), P. J. van Helsdingen (Leiden), L. B. Holthuis (Leiden), O. Karsholt (Copenha-
gen), O. A. Saether (Oslo), S. Shinonaga (Tokyo).
Dates of publication
Volume 133 (1), pages 1-120, 31 July 1990
Volume 133 (2), pages 121-280, i-viii, 14 December 1990
©
Nederlandse Entomologische Vereniging, Amsterdam ISSN 0040-7496
iv
Volume 133, no. I, 1999 AE
Ti üdschrift |
voor
| Entomologie
A journal of systematic and evolutionary
[ritomalazy since 1858
Published Bane Nederlandse Ento
Tijdschrift voor Entomologie
A journal of systematic and evolutionary entomology since 1858
Scope
The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a
long tradition in the publication of original papers on insect taxonomy and
systematics. The editors particularly invite papers on the insect fauna of the
Palaearctic and Indo-Australian regions, especially those including evolutionary
aspects e.g. phylogeny and biogeography, or ethology and ecology as far as
meaningful for insect taxonomy. Authors wishing to submit papers on disciplines
related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology
and applied entomology, are requested to contact the editorial board before
submitting. Usually, such papers will only be published when space allows.
Editors
E. J. van Nieukerken (elected 1986) and J. van Tol (1985)
Co-editors
A. W. M. Mol (1990) and R. T. A. Schouten (1990)
Advisory board
M. Brancucci (Basel), Y. Hirashima (Fukuoka) and M. R. Wilson (London).
The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the
‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society),
Amsterdam.
Editorial address
c/o National Museum of Natural History,
Postbus 9517, 2300 RA Leiden, The Netherlands.
Correspondence regarding membership of the society, subscriptions and
possibilities for exchange of this journal should be addressed to:
Nederlandse Entomologische Vereniging
c/o Instituut voor Taxonomische Zoölogie
Plantage Middenlaan 64
1018 DH Amsterdam
The Netherlands
Subscription price per volume Hfl. 300.- (postage included).
Special rate for members of the society. Please enquire.
Instructions to authors
May be obtained from the editors.
Graphic design
Ontwerpers B.V., Aad Derwort, 's-Gravenhage
Editorial
NINE te Jen) le A,
The editors of the ‘Tijdschrift voor Entomologie’ en the
first issue of volume 133, complete with a new cover design and a reshaped
graphic page design.
Both the cover and the lay-out were designed by Aad Derwort of the Dutch
firm ‘Ontwerpers’, The Hague. The cover depicts a variety of insects from a
collection of illustrations for graphic designers and illustrates the variety of topics
covered. The logo of the Netherlands Entomological Society can still be found
on the new cover, albeit somewhat smaller. This logo, in use since 1953, depicts
the Large copper (Lycaena dispar batavus Oberthür) on top of its greatly enlarged
egg. The logo was introduced on the cover of volume 96 when the late Alex
Diakonoff became the managing editor.
The lay-out of the articles has undergone relatively small changes compared
with previous issues.
ONTWERPERS
he
rchitectonische vormgeving
The new design parallels changes in the editorial policy of the ‘Tijdschrift’ to
transform it into an international journal of systematic and evolutionary ento-
mology. Although the journal has been published for an international audience
for several decades, the Dutch language was still used on the cover and elsewhere.
This has now all been translated into English, except the journal's title which has
become too well-known to be changed. Furthermore the editors now explicitly
express preference for articles on the insect fauna of the Palaearctic and Indo-
Australian regions. Papers on other regions are in principle only accepted if space
allows or if the paper deals with a topic of broader interest.
Some changes have gradually been introduced in the last three volumes. The
system of publishing a number of issues per year with one article each was left
in favour of two issues with various contents. This has reduced the expenses and
has given the opportunity to accept also short papers and book reviews. The
journal will usually be published around June Ist and December Ist. The editors
regret that the present issue appears some weeks late.
As can also be seen in this issue, the ‘Tijdschrift’ is certainly not only a medium
for Dutch entomologists or for members of the Netherlands Entomological
Society alone. Authors from all countries are cordially invited to submit their
papers. Only quality and consistency with the scope play a role in acceptance,
although we would certainly welcome new members of the society. It is general
policy now to invite external referees to review the manuscripts with regard to
their scientific quality and soundness. A new set of instructions for authors will
be published in the next issue and a guide for preparation of manuscripts on
diskette will be distributed on request.
The ‘Tijdschrift voor Entomologie’ has a long tradition and is certainly one of
the oldest entomological journals still being published. The journal has been
published annually since 1858, accounting for a total of 132 volumes up to now.
In the early days mainly the Dutch language was used. Later French, German and
English gradually started to play a role. In the second half of this century Dutch
lost its position as a scientific language and disappeared from the journal. From
the start the journal has especially served as a medium for systematic entomology
and, until the fifties, also for Dutch faunistics. Prior to 1953 it also included the
proceedings of the meetings of the Netherlands Entomological Society. Faunistic
papers, articles in the Dutch language and smaller papers of international interest
are now being published in 'Entomologische Berichten’ (Entomological Reports),
another society journal. Other publications of the Society are the international
journal "Entomologia experimentalis et applicata’ and the monograph series
‘Monografieën van de Nederlandse Entomologische Vereniging’.
The editors hope that the new face of the journal will please the readers and
especially that the contents, which is after all the most important part of a
journal, will continue to find their way to the professional and amateur entomol-
ogist.
Erik |. van Nieukerken
Jan van Tol
[editors]
A. W. M. Mol
R. T. A. Schouten
[co-editors]
A. ASQUITH & J. D. LATTIN
Systematic Entomology Laboratory, Department of Entomology, Oregon State University
NABICULA (LIMNONABIS) PROPINQUA (REUTER)
(HETEROPTERA: NABIDAE): DIMORPHISM,
PHYLOGENETIC RELATIONSHIPS AND BIOGEOGRAPHY
Asquith, A. & J. D. Lattin, 1990. Nabicula (Limnonabis) propingua (Reuter) (Heteroptera:
Nabidae): dimorphism, phylogenetic relationships and biogeography. - Tijdschrift voor
Entomologie 133: 3-16, figs. 1-15, tabs. 1-3. [ISSN 0040-7496]. Published 31 July 1990.
The taxonomy and distribution of the North American species Nabicula propinqua
(Reuter), is reviewed. The macropterous form occurs in both sexes, but is much less common
than the brachypterous form. Macropterous forms of both sexes have wider pronota. Males
have disproportionately longer antennae than females. Male and female genitalia of N.
propinqua are described and illustrated. The genitalia of individuals from isolated west coast
populations differ only slightly from those of eastern populations. A cladistic analysis of the
subgenus Limnonabis Kerzhner is presented and a new species-group classification sug-
gested. A biogeographic hypothesis explaining the present distribution of Limnonabis is
presented. We suggest that N. propingwa is a vicariant species that was isolated from a
population contiguous with that in western Europe with the opening of the Atlantic Ocean.
Correspondence: Adam Asquith, Systematic Entomology Laboratory, Department of Ento-
mology, Oregon State University, Corvallis, OR 97331, USA.
Key words. - Nabidae, sexual dimorphism, wing dimorphism, biogeography, phylogeny.
INTRODUCTION
Nabicula (Limnonabis) propinqua (Reuter), re-
cognized by its large size and elongate, slender
form, is one of the most conspicuous North Amer-
ican species of Nabidae (fig. 1). Unlike some of the
commonly encountered species of Nabis that occur
on vegetation in a variety of habitats, N. propinqua
is rare in collections and lives on the ground or low
vegetation in marshy habitats (Blatchley 1926). In
addition to N. propinqua, five other species in the
genus Nabicula Kirby occur in North America
(Henry & Lattin 1988). N. propinqua, however, is
the only member of the subgenus Limnonabis
Kerzhner in North America, the other five species
occur in Europe and Eastern Asia.
The most recent nomenclature (Kerzhner 1988)
considers Nabicula a subgenus of Nabis, and does
not recognize Limnonabis as a taxonomic group. In
this paper, however, we use the nomenclature of
Kerzhner (1981) because this classification has also
been used in recent faunistic works for Europe (Pé-
ricart 1987) and North America (Henry & Lattin
1988). In addition, species relationships within Na-
bicula are still unresolved and we anticipate the
nomenclature of Kerzhner (1988) to change again
(unpublished data).
Reuter (1872) described N. propinqua in Nabis
Latreille from a single brachypterous female spe-
cimen from Wisconsin, and described vicarius (in
Nabis) from a single brachypterous male from Il-
linois. Reuter (1880) synonymized vicarius with
propinqua. Hart (1907) described elongatus (in Na-
bis) from a single macropterous male from Havana,
Illinois. This species was synonymized with propin-
qua by Van Duzee (1916).
In this paper, we summarize the distribution and
habits of N. propingua and describe the internal
genitalia for both the male and female. We also
present a cladistic analysis of the subgenus Limno-
nabis and provide a hypothesis of the biogeographic
history of the taxon.
DESCRIPTIVE PART
Distribution
Nabicula propinqua occurs along the eastern sea-
board from Maryland to Maine and extends west-
ward between 40 and 50 degrees N latitude
throughout the Great Lakes region to North Da-
kota (fig. 2). West of the 100th meridian, N. pro-
pinqua occurs much farther north, with records
N
d
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
from Alberta, Manitoba, Quebec and above 60 de-
grees N latitude along Great Slave Lake in the
Northwest Territory. Essentially the distribution of
N. propinqua follows the Saline Lakes and Forest
Zone of Freshwater Lakes limnological regions of
Northcate & Larkin (1966). There are at least two
populations of N. propingua that are disjunct from
the main distribution. We have collected N. propin-
qua from two localities in the coastal marshes in
Oregon and have examined a single specimen from
Meade Co., Kansas.
With the exception of the latter three records,
most of the known distribution of N. propinqua
occurs within the area occupied by the ice sheet of
the Wisconsin Glaciation. If N. propinqua was
present in North America during the Pleistocene,
as we will argue, it must have been restricted to
areas south of the ice sheet or in one of the far
northern refugia (Matthews 1979). Because N. pro-
Fig. 1. Nabicula propingua (Reuter). Brachypterous male.
Dorsal habitus.
pinqua is not yet known to occur in the areas of
northern glacial refugia, it seems likely that N.
propinqua moved into the areas it now occupies
from areas south of the Wisconsin ice sheet some-
time within the last 10,000 years. In this scenario,
the Kansas specimen probably represents a Pleis-
tocene relictual population in an area previously
occupied by N. propinqua. Other such populations
are likely to be located (e.g. the marshes in the Sand
Hills of western Nebraska).
The origin of the Oregon coast population is
more perplexing. N. propingua has not yet been
found in the arid Inter-mountain region, nor in the
mesic Willamette Valley and Puget Trough areas of
Oregon and Washington. It appears to be restricted
to the coastal marshes of Oregon and possibly
Washington. Lattin (1966) suggested that this po-
pulation might be an introduction from the eastern
United States. Now, however, we believe this un-
likely, considering the restricted habitat of this spe-
cies and the fact that the Oregon population dis-
plays a slightly different genitalic structure than
eastern populations (see below).
If N. propinqua or its direct ancestor was present
in North America by the Eocene as we hypothesize
(see Discussion), there were few barriers to impede
Fast-West movement and a contiguous distribution
across the northern latitudes would have been pos-
sible. Beginning in the Miocene, orogenic activity in
western North America resulted in topographical
geographic barriers and, perhaps more impor-
tantly, increased aridity in the Intermountain re-
gion. Because N. propinqua lives in moist, riparian
habitats, this dramatic change in climate and terrain
would have restricted mobility and could have led
to the extinction of many, or all of the intervening
populations.
This is merely a hypothesis of course, and we
cannot actually date the separation of the west coast
population and it is possible that it is a much more
recent event. The climate of the northern United
States during the Pleistocene periods was much
more mesic than the present, with abundant, large
shallow lakes and marshes throughout the Great
Basin province (Smith 1978). This would have pro-
vided abundant habitats for N. propingua from the
Rocky Mountains west. With the advent of the
Hypsithermal and higher temperatures, these hab-
itats in this area largely disappeared (Barnosky et
al. 1987), which could have left the coastal popula-
tion isolated from those north and east of the Rock-
ies. This hypothesis would be corroborated if relic-
tual populations of N. propingua were found in
isolated marshes in the northern Great Basin.
Dimorphism
In many nabids, males and females often differ
markedly in size, occurrence of wings and propor-
ASQUITH & LATTIN: Nabicula propingua
? /
SCALE \ \
| °
o__ioo 200 300 400 500 600 700 800 900 1000 MILES IS
6” 200° 400 600 800 1000 1200 1400 KILOMETERS |
LAMBERT’S AZIMUTHAL EQUAL-AREA PROJECTION
©
\
| |
|
120
|
110
fl
90
x
WEST LONGITUDE
Fig. 2. Distribution of N. propingua in North America. Circles represent specimens examined, triangles are literature
records only. Solid line indicates maximum extent of Wisconsin ice sheet.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
2.2
PRONOTAL WIDTH (mm)
9 10 11 12 13
TOTAL LENGTH (mm)
Fig. 3. Relationship between the posterior width of the pronotum and total length in N. propinqua. y = 0.093239(x)
+ 0.465369, r? = 0.565. Open squares are brachypterous specimens. Solid squares are macropterous specimens.
29 TONDE
2.8 n
241
2.6
2.5
2.4
2.3
LENGTH ANTENNAL SEGMENT II (mm)
2.2 Wo Lia cs a la. AI
9 10 11 12 13
TOTAL LENGTH (mm)
Fig. 4. Relationship between the length of the second antennal segment and total length in N. propinqua. Open squares
are males. y = 0.13069(x) + 1.318405, r? = 0.149. Solid squares are females. y = 0.077957(x) + 1.62609, r? = 0.101.
6
ASQUITH & LATTIN: Nabicula propingua
Table 1. Comparison of six measurements among brachypterous and macropterous male and female Nabicula propinqua.
Data are mean and (range) in mm.
ALE. VERT PPW PL Al A2
male 9.8 0.405 1:39 152 1.66 2.60
(9.1-10.6) (0.38-0.44) (123-151) (1.41-1.69) (1.51-1.85) (2.28-2.86)
Brach M 10.5 0.41 1.97 1.69 1.70 2.69
female 11.8 0.444 1.57 1.73 1.65 DSS
(11.0-13.2) (0.39-0.48) (1.39-1.86) (1.61-1.86) (1.53-1.85) (2.36-2.82)
Brach F 11.8 0.41 2.09 1.86 1.60 2.49
(10.3-12.1) (0.40-0.42) (2.04-2.14) (1.83-1.89) (1.59-1.60) (2.48-2.50)
tions of body parts, but rarely are these differences
quantified. We examined sexual dimorphism in six
measurements, total length (TL), length of anten-
nal segment one (A1), length of antennal segment
two (A2), width of the vertex (VERT), posterior
pronotal width (PPW) and pronotal length (PL).
Measurements using an ocular micrometer were
made of 21 males and 32 females from throughout
the range of N. propinqua. We found differences
between sexes in four of the six measurements. TL,
VERT, PPW and PL, were greater in females than
in males, but there were no differences between
males and females for Al and A2 (table 1). The
differences between sexes seen in the former char-
acters are explained by the positive linear relation-
ship between these characters and the absolute size
of the individual. For example, PPW increases with
TL, and because females achieve a greater TL than
males, they also display a greater PPW (fig. 3). We
do not consider these characters true sexual dimor-
phisms therefore, but only size dependent charac-
ters.
The lengths of antennal segments also displayed
an increase with TL, but only within a given sex.
This is demonstrated by the fact that the slopes of
the A2-TL regressions for the two sexes are the
same (F = 0.415, P = 0.523), but the Y intercepts
are different (F = 8.662, P = 0.005) (fig. 4). This
indicates that although females are larger than
males (greater TL), small males have the same
length antennae as small females and large males
have the same length antennae as large females.
Thus, in males, the antennae are disproportion-
ately longer than in females. The length of the
antennae seems to be uncoupled from the general
correlation with body size that other measurements
show. This relationship does not appear to be com-
mon in the Nabidae. In a preliminary analysis of
four species of Nabis, males did not have dispropor-
tionately longer antennae, but rather there was a
linear relationship between antennal length and
size across the sexes.
Macroptery
Nabicula propinqua usually occurs in the bra-
chypterous state, with the apex of the fore wings
reaching to the middle of the second abdominal ter-
gite (fig. 1). Macropterous females have been re-
ported by Harris (1928) and Froeschner (1971). On-
ly 2 of 45 females we examined were macropterous.
In these individuals, the fore wings reached the mid-
dle of the seventh abdominal tergite. The hind wings
are equally well developed, reaching to the anterior
margin of the seventh abdominal tergite (fig. Sa).
Fig. 5. N. propinqua A. Macropterous female. B. Macrop-
terous male.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Fig. 6. Seminal depository of female N. propinqua. Dorsal
view. A. Sclerotized ring. B. Membranous sac. C. Muscular
lobe ventral to oviducts.
We have examined 29 male specimens and have
seen only one macropterous individual (fig. 5b).
This specimen was the holotype of Nabis elongatus
Hart and is deposited in the Illinois Natural History
Survey collection (INHS). Although Harris exam-
ined the material in the INHS (we have seen spe-
cimens with his determinations), he apparently
overlooked this specimen, because he stated (Harris
1928) that males may never occur in the macrop-
terous state. In England and on the Continent, only
the females of Nabicula lineata (Dahlbom) are
known in the macropterous form (Southwood and
Leston 1959). Macropterous females and males
have been reported for N. ussuriensis (Kerzhner)
in eastern Asia (Kerzhner 1968).
Differences in the structure of the pronotum
between brachypterous and macropterous speci-
mens have been reported for both the latter two
species. In N. /ineata, the pronota of brachypterous
specimens are 1.1 times as long as wide and in the
macropterous forms the length and width are equal
(Pericart 1987). In N. wssurtensis, the pronotum in
brachypterous specimens is 1.1-1.2 times as long as
wide and only 0.94 times as long as wide in macrop-
terous specimens (Kerzhner 1968).
In N. propinqua, the ratio of PL to PPW is lower
in the macropterous form (0.85-0.89) than the bra-
chypterous form (1.05-1.15) in both males and fe-
males. The lower ratio results not from a shorten-
ing of the pronotum in the macropterous form, but
from an increase in the width (fig. 3). In particular,
the posterior lobe of the pronotum is greatly flared.
8
The increased width of the pronotum is probably an
indirect effect of the development of flight muscles
and phragmata in the pterothorax of the winged
form (Darnhoffer-Demar 1969). Because the ante-
rior end of the mesothorax is enclosed by the pos-
terolateral angles of the pronotum, an increase in
diameter of the mesothorax would cause a similar
change in the posterior lobe of the pronotum.
Female Genitalia
The seminal depository is large and dome-
shaped, consisting of a fleshy external portion and
a partially sclerotized internal structure, the apex of
which bears three to five blunt sclerotized teeth (fig.
6). A thick muscular structure lies between the base
of the depository and the oviduct. The sclerotized
ring is a single structure occupying the ventral sur-
face of the left side of the depository, extending
from the depository laterally and curving ante-
riorly. It is surrounded by a large membrane also
directed anteriorly.
Male Genitalia
Parameres
Reuter (1872) and Harris (1928) provided lateral
views of the left paramere of Nabicula propingua.
Paramere morphology in some groups of Nabidae
is very conservative (e.g. the genus Nabis) and N.
propingua displays this general form (fig. 7).
Within the subgenus Limnonabis, the paramere of
N. propinqua most closely resembles that of N.
pontica. In contrast, N. lineata displays an unusually
elongate apex. In some specimens of N. propingua,
the apex of the paramere is slightly bent laterally,
a condition which is strongly developed in N. ussu-
riensis (Kerzhner) and N. demisa (Kerzhner 1968,
1981).
b
Fig. 7. Right paramere of N. propingua A. Lateral view
B. Medial view. C. Ventral view.
Fig. 8. Male aedeagus of N. propinqua from Oregon.
Sclerites indicated by roman numerals. IVa. sclerite IV of
specimen from Illinois.
Aedeagus
The aedeagus of Nabicula propinqua is very sim-
ilar to that of N. lireata and N. pontica. The mem-
branous portion of the aedeagus has four sclerites
(fig.8). No nomenclature has been proposed for the
aedeagal sclerites of the Nabidae, and here we
number the sclerites with Roman numerals begin-
ning at the base of the aedeagus (sclerite I) and
moving to the apex (sclerite IV). Sclerite I is linear
and parallel to the longitudinal axis of aedeagus; it
has a wide base and slightly tapering distalarm with
a blunt, rounded apex. Some specimens may display
a slight flange on this sclerite. Sclerite II is pistol-
shaped, with a wide, arcuate base, a curved obtuse
angle near its midpoint and a tapering, rounded
distal arm. The distal arm bears flanges on both
edges reaching from angle to apex. Sclerite III is
roughly quadrangular and distinctly comb-shaped,
with five short, blunt teeth directed toward the
midline of the aedeagus. Sclerite IV is situated
transversely in the aedeagus, with a broad, oval
base. This sclerite exhibits geographic variation. In
specimens from the Oregon coast, the sclerite has
a thick, bluntly rounded arm (fig. 8 IV), and in
specimens from east of the Rocky Mountains, the
arm is longer, thinner, with a more pointed apex.
(Fig. 8 IVa). There are two areas with sclerotized
ASQUITH & LATTIN: Nabicula propinqua
denticles along the folds of the membrane, one area
distal to sclerite II and the other distal to sclerite IV.
These denticulate areas are also present in identical
form in related species.
PHYLOGENETIC ANALYSIS
To understand the origin and evolution of Nabz-
cula propinqua in North America, it is necessary to
identify its phylogenetic relationship to other taxa.
Within Nabicula, Kerzhner (1981) recognized
three subgenera, Limnonabis Kerzhner, Dolicho-
nabis Reuter, and Nabicula Kirby. He distinguished
the subgenus Limnonabis from Dolichonabis
(sensu strictu) by the diverging posterior lobe of the
head, recurved connexivum of the male and the
presence of three or four sclerites in the aedeagus.
Within Limnonabis, he identified two groups, the
lineata group composed of N. lineata, N. pontica
and N. propingua, identified by the large parameres
and multiple rows of spines of Ekblom’s organ (see
fig. 11). The wsswriensis group contains N. ussw-
riensis (Kerzhner), N. demisa (Kerzhner) and N.
sauteri (Poppius), united by the small parameres
and the single row of spines (see fig. 10).
To further clarify the relationships among these
taxa, we conducted a cladistic analysis of the subge-
nus Limnonabis using the computer program
HENNIG 86 (Farris 1988). Because we were not able
to examine specimens of all species in the subgenus,
we used characters that were described and/or illus-
trated for the other taxa by Kerzhner (1968, 1981)
and Péricart (1987). We used structures of the male
Anaptus
flavomarginata
limbata
sauteri
demisa
ussuriensis
propinqua
lineata
— pontica
Limnonabis
Fig. 9. Cladogram of Limnonabis species based on table
2. Nodes denoted by letters in circles. Characters and
character states of synapomorphies given under each
node.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
10
ines.
, multiple rows of sp
. propinqua
BEN
ines
le row of spi
, sing
Ekblom's organ. 10. N. americolimbata
ing
iew of male genital capsule show
10L V
Figs. 10-11. Poster
Figs. 12-13. Structure of individual spines of Ekblom's organ. 12. N. americolimbata. 13. N. propinqua.
ASQUITH & LATTIN: Nabicula propinqua
Table 2. Description of characters used in the cladistic analysis of the subgenus Limnonabis. Character states are preceeded
by their coding in the data matrix and (in parentheses) the number of times state arises in the cladogram.
Female Genitalia
1. 0(1) Double sclerotized ring.
1(1) Single sclerotized rings.
2. 0(1) Sclerotized ring symmetric, located medio-
dorsal.
1(1) Sclerotized ring asymmetric, located laterally.
3. 0(1) Membranous “sac” around sclerotized ring
absent.
1(1) Membranous "sac symmetric, located dor-
sally.
2(1) Membranous sac” asymmetric, not located
dorsally.
4. 0(1) Muscular lobe” antero-ventral to sperm
ducts absent.
1(1) Muscular “lobe” antero-ventral to sperm
ducts present.
Male Aedeagus
5. 0(1) Aedeagus with fewer than three sclerites.
1(1) Aedeagus with 3 sclerites.
2(1) Aedeagus with 4 sclerites.
3(1) Aedeagus with more than 4 sclerites.
6. O(1) Length of sclerite I less than four times its
width at middle.
1(1) Length of sclerite I at least four times its
width.
7. 0(2) Sclerite I without flanges present.
1(1) Sclerite I with flanges.
2(1) Sclerite I with flanges only slightly deve-
loped.
8. 0(1) Sclerite II absent.
1(1) Length of sclerite II less than three times its
width.
2(1) Length of sclerite II three times its width.
paramere (2 characters), aedeagus (6 characters),
male genital capsule (1 character), female genitalia
(3 characters), body structure (4 characters) (table
2). Determining the polarity of some of the char-
acters was difficult, because the homology of some
of the structures in other genera could not be deter-
mined and no previous cladistic analysis within the
Nabidae was available for comparison. Because of
these problems, we included the following three
taxa in our analysis as outgroups, Nabicula
(Dolichonabis) limbata (Dahlbom); Nabicula
(Nabicula) flavomarginata (Scholtz); Anaptus ma-
jor (A. Costa) (table 3). These taxa were chosen
because they represent the other two subgenera of
Nabicula and a more distant member of the tribe
Nabini. All multistate characters were coded as or-
dered except characters 5 and 10 because we were
uncertain of their transformation sequences.
The analysis produced a single tree of minimal
length (31 steps, consistency index of 83.0, fig. 9).
The subgenus Limnonabis appears to be a mono-
phyletic group, as indicated by component E of the
cladogram. It is identified by characters 14 and 15,
the diverging posterior margin of the head, and the
recurved connexivum of the male respectively.
9. 0(1) Sclerite II without flanges.
1(1) Sclerite II with flanges.
10. 0(1) Sclerite III linear without flanges.
1(1) Sclerite III prong-shaped.
2(1) Sclerite III comb-shaped.
3(1) Sclerite III short, not linear.
4(1) Sclerite III linear with flanges.
11. 0(1) Sclerite IV absent.
1(1) Sclerite IV with recurved arm.
2(1) Sclerite IV comb-shaped.
Male Paramere
12. 0(1) Tip of paramere not elongate.
1(2) Tip of paramere elongate.
13. 0(1) Apex of paramere straight.
1(2) Apex of paramere bent laterally.
External Structure and Form
14. 0(1) Head straight or converging behind eyes.
1(1) Head diverging behind eyes.
15. 0(1) Medial half of male connexivum straight.
1(1) Medial half of male connexivum curved un-
der and appressed to abdomen.
16. 0(1) Spines of Ekblom’s organ in a single linear
row.
1(1) Spines of Ekblom's organ in a bunched row.
17. 0(1) Body length less than 5 times the width.
1(1) Body length greater than 5 times the width.
These are two of the characters Kerzhner (1968)
used to define the subgenus. We are doubtful of the
integrity of the first character, the diverging poster-
ior lobe of the head. We have examined species in
other genera (Nabis Latreille, Lastomerus Reuter)
in which this character displays almost the same
development as seen in Limnonabis. At this time,
we can find no genitalic characters that unite all
species currently placed in Limnonabis and the only
character that we feel defines the group is the re-
curved connexivum of the male.
The wssuriensis group does not appear to be a
natural one, its members are united only by plesi-
omorphic characters, such as the single row of
spines of Ekblom’s organ. Nabicula ussuriensis is
actually united with the /neata group by the pres-
ence of an asymmetric sclerotized ring (2-1) (char-
acter-character state), presence of a membranous
“sac! around sclerotized ring (3), aedeagus with
four sclerites (5-2), length of sclerite II three times
its width (8-2) and sclerite II with flanges (9-1)
(component C). The /neata group of Kerzhner
(1968) is clearly monophyletic, represented by com-
ponent B on the cladogram and defined by the
asymmetric membranous sac (3-2), muscular lobe
Il
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 3. Character matrix for Limnonabis processed by HENNIG86.
l 2 5) 4 5 6 7 8 9) JO) iil 2 15 14 15 16.17
Anaptus t © @ @ SOON cane = JO) te 122 ? 1 © © © © 0
flavomarginata O © © © OA EE QE ? 2 © i 0 © © @ ©
limbata l © © 0 Od. 1 lame? ? CE) LI ® © © © »@
sauteri Lr @ © 8 1 1 TOO ANO Om 200 3] Io 30
demisa L @ Orr © ROME Oso ORT er Od 1 TO OS OI
usseriensis 1 1 le 2 1 1 2 dm ERD 0 1 1 LAON, dl
propinqua 1 1 2 1 2 an 2 1 2 1 @ O 1 1 1
lineata 1 1 2 l 2 1 0 2 Il 0 1 1 0 l Il Il Il
pontica 1 le 2 1 2 OZ OR I ® @ A 1 1 1
behind the sperm ducts (4-1), and the bunched row
of spines comprising the Ekblom’s organ (16-1).
The latter character appears to be unique among
the Nabidae. Nabicula propinqua, N. lineata, and
N. pontica display a bunched row, 3-5 spines wide,
on each side of the anal tube (fig. 11). All other
species of nabids that we have examined possess
only a single row of linearly arranged spines (fig.
10). The structure of the individual spines may also
prove to be an informative character. In N. propin-
qua, the spines are thin, with the distal third sharply
narrowed, sinuous, with the apex curved laterally
(fig. 13), while in N. (Limnonabis) americolimbata
the spines are wider, with the distal halves flattened
and expanded (fig. 12).
Nabicula lineata and N. pontica are united by
having sclerite III linear and without flanges (10-0)
and sclerite I without flanges (7-0). This latter char-
acter is homoplasious, because the 0 state also ap-
pears in N. demisa.
BIOGEOGRAPHY
All members of the subgenus Limnonabis, with
the exception of Nabicula propinqua, are found in
the Palaearctic region. The three species arising
i i SCHOOL SERIES
Outline Map
WORLD
- Scale of Milen at Equator
» iP A 1000 2000 3000
MAP NO. 1739
+ + — del
| SC — | COPYRIGHT *
AMERICAN MAP CO., INC.
WORLD MAP CO., INC.
mn
R 1
100, 120, 160, 10 180 160 140 120 100
Fig. 14. Distribution of the subgenus Limnonabis. Vertical lines and solid circles in North America, N. propingua. Large
dots in Europe, N. lineata. Small dots in southeastern Europe, N. pontica. Slanted lines with small dots in southeast
Asia, composite ranges of N. ussuriensis, N. demisa and N. sauteri.
12
lineata (Europe)
pontica (Europe)
Eocene) ——————
propinqua (N Am)
LR, ———
Cretaceous
ussuriensis (NE Asia)
Fig. 15. Cladogram of Limnonabis with the present dis-
tribution indicated after each species name. Hypothes-
ized ages of intercontinental vicariances indicated at left.
from the cladogram first, N. ussuriensis, N. demisa
and N. sauteri occur in far eastern Asia (fig. 14).
Two members of the /ineata group, N. lineata and
N. pontica occur in Europe and the Mediterranean
region and N. propingua occurs only in North
America. With regard to the views of Ball (1975),
we generated a testable biogeographic hypothesis
to explain the present distribution of the members
of the monophyletic group represented by compo-
nent C on the cladogram. We chose to use only this
group and not all of Limnonabis, because N. sauteri
and N. demisa are primitive taxa whose placement
in the subgenus appears tenuous.
We hypothesize that the distributions of the spe-
cies in component C are the result of vicariance
caused by the breakup of Laurasia beginning in the
Cretaceous. With the development of the Turgai
straits in the late Cretaceous, Asiamerica and Eur-
america were separated into two distinct land
masses (Cox 1974). This would have resulted in the
ancestral population being divided into two groups,
with the ancestor of N. ussuriensis in western Asi-
america and the ancestors of the /ineata group in
Euramerica.
The ancestor of the /ineata group was distributed
throughout Euramerica and with the opening of the
North Atlantic eventually separating North Amer-
ica from Europe in the Eocene, the ancestor of N.
propinqua was isolated from that of N. /imeata and
N. pontica. The exact time of separation of these
two groups cannot be known because there were
several distinct land bridges that closed at different
times, with at least one dispersal route across the
Thulean bridge which may have persisted into the
Miocene 20 millions years ago (Noonan 1988).
With the exception of the elongate apex of the
paramere in N. lineata, N. lineata and N. pontica
are very similar, suggesting that the separation of
these two species is a more recent event.
In summary, we invoke two vicariant events to
ASQUITH & LATTIN: Nabicula propinqua
explain the origin and present distribution of the
taxa in component C of the cladogram (fig. 15).
1) The ancestor of N. ussuriensis was isolated in
Asiamerica from component B in Euramerica by
the Turgai Straits. 2) The ancestor of N. propinqua
was isolated from component A in Europe by the
opening of the Atlantic. If our phylogenetic hy-
pothesis is correct, we believe this biogeographic
scenario to be the most parsimonious one. A disper-
sal hypothesis for the evolution of component C
would require the ancestral form to have moved
into, and then gone extinct from an area reaching
from Eastern Europe to Eastern Asia. Likewise,
dispersal of the ancestor of N. propinqua from
Europe to North America via Beringia as suggested
by Schaefer & Calabrese (1980), would require the
extinction of this taxon in all of northern Asia.
Another hypothesis for propinqua reaching North
America is by dispersal across the North Atlantic.
Because of the specialized habits, low incidence of
macroptery and low vagility of this species (it has
not been recorded from flight traps or aerial sam-
pling), we consider the latter hypothesis unlikely.
DISCUSSION
Using examples from the Trichoptera, Homop-
tera and Coleoptera, Allen (1983) discussed the
North America - Europe - Northeast Asia distri-
butions for insects. He found a common pattern
among these groups in which the North American
and European taxa shared a common ancestor after
the origin of the Northeast Asian groups. Similar
to our hypothesis for Limnonabis, Platnick (1976)
discussed the vicariant patterns in the spider genus
Callilepis (Gnaphosidae). His phylogenetic analysis
of the genus showed that species groups were dis-
tributed in areas that reflected the breakup of Laur-
asia. He also suggested that the Turgai straights
separated a Asiamerica group from a Euramerica
one, and the Atlantic rift further separated the
Euramerica group.
The species in the N. /ineata group are very
similar morphologically, differing primarily in the
structure of the internal genitalia. It may seem un-
usual that we invoke such great ages for the sepa-
ration of these species in light of the slight degree
of morphological divergence. Without rehashing
the old and continuing debate regarding rates of
evolution, we feel that our hypothesis is tenable for
three reasons. First, with few exceptions (e.g. Na-
bicula subcoleoptrata Kirby) the lineages within the
tribe Nabini appear to be highly conservative or
canalized in their morphology. Many species differ
only slightly in external morphology and are distin-
guished primarily by the male aedeagus and the
seminal depository of the female.
Second, explosive speciation and morphological
evolution is often correlated with changing envi-
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
ronments or lineages radiating into new adaptive
landscapes (Simpson 1944), while groups occupy-
ing stable, constant and predictable environments
may undergo very little change through time. The
lineata group inhabits moist, marshy habitats and
secondarily, estuarine marshes, environments that
prevailed during the Cenozoic but also have per-
sisted relatively unchanged to the present. Consid-
ering the stability of morphology in the group, low
vagility and their specific, unchanged habitat, large
scale vicariant events resulting in superficially sim-
ilar, disjunct species might be expected.
Finally, it is becoming increasingly apparent that
in some groups of insects, extant species are of great
antiquity. In the Coleoptera for example, the Pleis-
tocene climatic oscillations greatly altered distribu-
tions but did not result in speciation (Coope 1970,
Matthews 1977). Some species of beetles are appar-
ently in excess of 10 million years old (Larsson
1978). Similar examples are available for the Heter-
optera, (Calabrese 1978, 1980).
ACKNOWLEDGMENTS
We thank Michael D. Schwartz for critically read-
ing this manuscript and for help in the data analysis
and Bonnie B. Hall for the fine habitus illustrations.
We thank the following for their assistance in ex-
amining critical specimens: Thomas J. Henry, Uni-
ted States National Museum of Natural History
(USNM); Randall T. Schuh, American Museum of
Natural History (AMNH); Kathryn C. McGiffen,
Illinois Natural History Survey (INHS); Charles A.
Triplehorn, Ohio State University; Philip J.
Clausen, University of Minnesota (UM); B.J. Har-
rington and S. Krauth, University of Wisconsin-
Madison (UWIM); Robert W. Brooks, University
of Kansas (KU); John A. Chemsak, University of
California, Berkeley (UCB); Paul H. Arnaud Jr.,
California Academy of Sciences (CAS); James Lieb-
herr and E. Richard Hoebeke, Cornell University
(CU); Frank Mead, Florida State Collection of Ar-
thropods (FSCA); Jeffrey C. Burne, University of
Wyoming; Edward U. Balsbaugh Jr., North Dakota
State University; Brett C. Ratcliffe, University of
Nebraska.
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Michigan. - Occasional Papers of the Museum of Zool-
ogy, University of Michigan. 118: 1-39.
Kerzhner, I. M., 1968. New and little know Palearctic bugs
of the family Nabidae (Heteroptera). - Entomological
Review 47: 848-863.
Kerzhner, I. M., 1981. Fauna SSSR. Nasekomye khobot-
nye, t. XIII, vyp. 2: Poluzhestkorylye semejtstva Nabi-
dae [Hemiptera of the family Nabidae]. - Leningrad,
326 pp.
Kerzhner, I. M., 1988. Family Nabidae. Ir: PH. Ler (ed).
Keys to the identification of insects of the Soviet Far
East. Vol 2: Homoptera and Heteroptera. Nauka. Le-
ningrad.
Larsson, S. G., 1978. Baltic Amber — a palaeobiological
study. - Entomonograph 1. Scandinavian Science Press
Ltd., Klampenborg. Denmark. 192 pp.
Lattin, J. D., 1966. Stalta-major (Costa) in North America
(Hemiptera: Nabidae). - Proceedings of the Entomo-
logical Society of Washington 68: 314-318.
Matthews, J. V. Jr., 1977. Tertiary Coleoptera fossils from
the North American Arctic. - Coleopterist's Bulletin
31: 297-308.
Matthews, J. V. Jr., 1979. Tertiary and Quaternary envir-
onments: historical background for an analysis of the
Canadian insect fauna. - Ir: H. V. Danks, (ed.), Canada
and its insect Fauna. - Memoirs of the Entomological
Society of Canada 108: 31-86.
Noonan, G. R., 1988. Faunal relationships between east-
ern North America and Europe as shown by insects. —
In: J. A. Downes & D. H. Kavanaugh (eds.). Origins
of the North American Insect Fauna. - Memoirs of the
Entomological Society of Canada 144: 39-53.
Northcate, T. G. & P. A. Larkin, 1966. Western Canada. pp.
451-485. — In: D. G. Frey (Ed.). Limnology of North
America. - University of Wisconsin Press. Milwaukee
& London.
Péricart, J., 1987. Hemipteres Nabidae: d'Europe Occiden-
tale et du Maghreb. - Faune de France 71: 1-185.
Platnick, N. I., 1976. Drifting spiders or continents? Vic-
ariance biogeography of the spider subfamily Larionii-
nae (Araneae, Gnaphosidae). - Systematic Zoology 27:
101-109.
Reuter, O. M., 1872. Nabidae novae et minus cognitae.
Bidrag till Nabidernas kannedom. - Ofversigt af Kon-
gliga Svenska Vetenskaps -Akademiens Förhändlingar
29(6): 79-96.
Reuter, O. M., 1880. Anteckninger om Coriscus lineatus
Dahlb. - Ofversigt af Finska Vetenskaps-Societatens
Förhändlingar 22: 25-32.
Schaefer, C. W. & D. M. Calabrese, 1980. Amphi-Atlantic
species-pairs in two genera of water striders (Hemi-
ptera, Gerridae). - Entomologia Generalis 6: 271-280.
Simpson, G. G., 1944. Tempo and Mode in Evolution. —
Columbia University Press. New York. 237 pp.
Smith, G. R., 1978. Biogeography of Intermountain fishes.
- In: K. T. Harper & J. L. Reveal (Eds.), Intermountain
biogeography: A symposium. - Great Basin Naturalist
Memoirs 2 :17-42.
Southwood, T. R. E. & D. Leston, 1959. Land and water
bugs of the British Isles. - Frederick Warne and Co.
Ltd., London and New York. 436 pp.
Strickland, E. H., 1953. An annotated list of the Hemi-
ptera of Alberta. — Canadian Entomologist 85: 193-
214.
Van Duzee, E. P., 1916. Chech List of the Hemiptera
(Excepting the Aphididae, Aleurodidae and Coccidae)
of America, North of Mexico. - New York Entomolo-
gical Society, New York, 111 pp.
Received: 18 Seprember 1989
Accepted: 1 March 1990
ASQUITH & LATTIN: Nabicula propinqua
APPENDIX
Literature Records
Canada. — Alberta: Edmonton (Strickland 1953).
United States. — Illinois: Cook Co.: Argo; West Pull-
man (Blatchley 1926). - Iowa: Hancock Co.: 8 mi. SE
Britt, July 6, 1928 (Hendrickson 1930). — Maryland:
Somerset Co.: Deal Island, June 29, 1970 (Froeschner
1970). - Michigan: Berrien Co.: New Buffalo & Stevens-
ville (Hussey 1922b). - New Jersey: Morris Co.: Snake
Hill; 2? Co.: White's Bay, July 20, 1914 (Froeschner
1971). - North Dakota: Ramsey Co.: Devils Lake, July 22
& 25 (Hussey 1922a). - South Dakota: Day Co.: Waubay,
August 21, 1924 (Harris 1943); Grant Co.: Bigstone,
August 20, 1924 (Harris 1943).
Specimens Examined
Canada. — Alberta: High Prairie, July 17, 1961, A. R.
Brooks, 1 6, 1 9 (OSU); Slave Lake, August 14, 1924, O.
Bryant, 2 4, 1 9 (CAS). - Manitoba: Red Deer R., August
3, 1937, C. L. Johnston, 1 & (KU); The Pas, August 11,
1937, D. G. Denning, 1 9 (UM). - North West Territory:
5 mi. SE Ft. Providence, August 15, 1965, J. & W. Ivie, 1
2 (AMNH). - Ontario: Thessolon Lake, shore, July 21,
1965, J. & W. Ivie, 19 (AMNH). - Quebec: Quinze Lake,
August 15, 1907, W.J. Palmer, 1 Q (CAS); Saskatchewan:
Qu’ Appelle River, N Tuxford, July 29, 1965, J. & W. Ivie,
292 (AMNH).
United States. — Illinois: Boone Co.: Belvidere, June
27, 1955, J.A. Slater (OSU); Fulton Co.: Havana, June 9,
1905 / Ill. Sands, Hart Coll. / Nabis elongatus Hart, type
/ Reduviolus elongatus Hart / TYPE Nabis elongatus, C.
A. Hart, 1 & (INHS); Lake Co.: Fox Lake, August 23,
1944, Frison & Ross, 4 6,9 2 (INHS); Lake Villa, swamp,
August 10, 1906 / Reduvolius vicarius Reuter, micr. F /
Nabis propinquus Reuter, H. M. Harris / Van Duzee Cat.
No. 825 Det. Harris, 1 9 (INHS); Waukegan, May 14,
1930 4 & (INHS); Waukegan, beach, August 23, 1906 /
Reduvolius vicarius Reut, Micro. M / Nabis propinquus
Reuter, Det. H. M. Harris / Van Duzee Cat. No. 825, Det.
Harris 1 @, | nymph (INHS). - Kansas: Meade Co:
September 13, 1944, R. H. Beamer, 1 & (OSU). - Maine:
Sagadahoc Co.: Popham Beach, September 4, 1920, A. P.
Morse, 1 9, (AMNH). - Massachusetts: Essex Co.: Beach
Bluff, 22 June, 1914, H. M. Parshley, 1 nymph (CAS); 13
August, 1914, ex. Carex, 2 8,4 9; June 21, 1915, 1 nymph
(CAS); August 17, 1916, ex. Carex, 3 6, 11 9 (AMNH);
Middlesex Co.: Faneuil, August 2, 1904, A. P. Morse, 1 6;
August 12, 1904, A. P. Morse, 1 9; October 1, 1904, A. P.
Morse, 1 Q (AMNH). - Michigan: Cheboygan Co.: July
25, 1940, L. Spencer, 1 Q (OSU); July 1, 1950 J. D. Lattin,
I nymph (OSU); Duncan Bay, July 27, 1957, W. J. Han-
gan, 2 9 (KU); Emmet Co.: August 10, 1950, J. D. Lattin,
1 2 (OSU); Huron Co.: Sand Point, June 24, 1922, R. 9.
Hussey, 1 9 (FSCA). - Minnesota: Saint Louis Co.: Ea-
glesnest, July 15, 1959, W. V. Balduf, 2 &, 1 9 (UM);
Traverse Co.: Lake Traverse, 7.5 mi SW Wheaton, July 27,
1974, B. Tollefson, 1 4, 1 9 (UCB). - New York: Catta-
raugus Co.: South Dayton, 23 July, 1946, R. H. Beamer,
1 Q (KU); Erie Co.: Buffalo, July 16, 1901, 1 9 (CAS);
Nassau Co.: Long Island, Cold Spring Harbor Biol. Lab.,
B. Darnall, 1 ® (CAS); Piermont, June 17, 1934, Scholt,
1 9 (AMNH). - North Dakota: Ramsey Co.: Devil's
Lake, July 22, 1920, T. H. Hubbell, 1 @ (FSCA). - Ohio:
Erie Co.: Cedar Point, Sandusky, August 31, 1905, Van
Duzee, 1 2 (UCB); Union Co.??, Camp Perry, September
5, 1921, W. L. McAtee, 1 Q (USNM). - Oregon: Coos Co.:
15
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
South Slough Sanctuary, SW Coos Bay, September 24.
1988, A. Asquith, 1 9 (OSU); Tillamook Co.: Island
Camp, near woods, July 18, 1959, K. Fender, 1 6 (OSU),
5 mi E Pacific City, meadow, September 9, 1962, J. Ca-
pizzi, 2 Q (OSU); Sand Beach St. Pk., Sand Lake, Sep-
tember 7, 1988, A. Asquith & J. D. Lattin, 4 4, 12 9
(OSU). - Wisconsin: Dane Co: TON, RTE, S28, stream
edge, aquatic net, September 14, 1973, J. Hender, 1 Q
(UWiM); Wood Co.?, Nevin Marsh, July 23, 1974, site 6,
sweep net, D. Bach, 1 @ (UWiM); July 11, 1974, site 2,
D. Bach, 1 6 (UWiM).
16
A. BEDOS & L. DEHARVENG
Université Paul Sabatier, Toulouse, France
NEW SPECIES OF SUPERODONTELLA STACH
(COLLEMBOLA: ODONTELLIDAE) FROM THAILAND
Bedos, A. & L. Deharveng, 1990. New species of Superodontella Stach (Collembola: Odon-
tellidae) from Thailand. - Tijdschrift voor Entomologie 133: 17-26, figs. 1-28, tabs. 1-3 [ISSN
0040-7496]. Published 31 July 1990.
The genus Superodontella Stach (Odontellidae) is recorded from Thailand for the first time.
Four new species are described, S. cicoria sp. n., S. gouzet sp. n., S. longispina sp. n. and
S. flammata sp. n. Some new morphological characters are presented.
A. Bedos and L. Deharveng, UA333 du CNRS, Laboratoire de Zoologie, Ecologie des
Invertébrés Terrestres, Université Paul Sabatier, 118 route de Narbonne, 31062 Toulouse
Cedex, France.
Key words. - Collembola, new species, Thailand.
INTRODUCTION
Superodontella was created by Stach in 1949 for
the species Odontella ewingi Folsom, 1916. Long
considered as a synonym of Odontella in the liter-
ature, it was redefined as a subgenus of Odontella
by Deharveng (1981a). It differs from Odontella s.
str. in the absence of an apical exsertile bulb on
antennal segment IV. This character has proved to
be consistent with biogeographical data, as no spe-
cies of Odontella s. str. has been found outside the
austral region in spite of extensive recent collecting
in Madagascar, tropical America and South East
Asia. Here we accept Superodontella as valid at
generic level with the diagnosis given by Dehar-
veng (1981a), because of the large number of spe-
cies now known to have this antennal character.
TAXONOMIC CHARACTERS USED IN THE
DESCRIPTIONS
The most important diagnostic characters for
Superodontella occur in the antennal and buccal
areas. Several new characters are introduced here.
Body length. — Adults are scarce, sometimes
absent in many Superodontella species. ‘Large juve-
niles’ are specimens devoid of any sexual differen-
tiation, but of a size as large as or larger than adults.
The biological problem underlying these observa-
tions is under study.
Integument granulations. — (1) Shape, size and
arrangement of secondary granules, particularly on
head and abd. VI (abdominal segment VI). (2) Ar-
rangement and morphology of muscular insertions
on head (which could be studied on other areas of
the body).
Antennae. — (1) Apical structures of ant. IV
(antennal segment IV). (2) Number and morphol-
ogy of s-setae on ant. IV. (3) Number of trumpet-
setae on ant. IV (mostly ventral; low intraspecific
variability in adult, but character instar-dependent).
(4) Form and length of s-setae on ant. III. (5)
Number of ordinary setae on ant. III.
Ocular area. — (1) Morphology of PAO (postan-
tennal organ) (‘amiboid’ with coalescence of lobes
at base, or ‘hypogastruroid’ with well separated
lobes). (2) Integument swellings between eyes.
Buccal area. — (1) Length and morphology of
buccal cone (range from very short and thick to very
long and thin). (2) Labrum chaetotaxy, difficult to
observe, is usually not mentioned in descriptions. A
conspicuous pair of longitudinal sclerotized struc-
tures terminating distally could be special modified
setae; they are present in all studied species (fig.
12). (3) Labium chaetotaxy, following Massoud
(1967) nomenclature (fig. 12). (4) Number and
length of perilabial setae (following the nomencla-
ture of Yosii (1971)) and postlabial setae (following
in part the nomenclature of Da Gama (1988) (fig.
19). Both nomenclatures are incomplete when all
Poduromorpha are considered; a general complete
nomenclature is presently under study. (5) Form
and length of maxilla stipa and fulcrum, including
those of E.P.P. (external posterior processus) and
LP.P. (internal posterior processus), highly discrim-
inant interspecific characters (fig. 6). (6) Number,
form and length of mandibles.
Dorsal chaetotaxy. — (1) Morphology of ordi-
nary setae, particularly on abd. VI. (2) Relative
length of s-setae. (3) Position of s-setae on the
tergites (stable in the species examined here). (4)
Presence/absence of c2 seta on head. (5) Setae of
central area (d- and sd-setae) are the same in all our
17
TIJDSCHRIFT VOOR ENTOMOLOGIE. VOLUME 133. 1990
/ à
\ IN, u
a8 4 i
È it r i "i
Figs. 1-7. Superodontella ciconia sp. n. — 1. Abdomen VI tergite; 2. Central area.of head between dl and d5 setae:
integument secondary granules and muscular insertions; 3. Right fourth antennal article, ventral side; 4. Right fourth
antennal article and distal part of ant. III, dorsal side; i: i-seta; ms: s-microchaeta; or: distal organite; s1 to s4: s-setae
of ant. II] organite; x: ordinary distal seta belonging probably to ant. III; 5. labial, perilabial and postlabial setae; 6. Maxilla
and mandible, dorsal view; fulcrum with arm, I.P.P. and E.P.P. (cf text); md: mandible; st: stipa; 7. Dorsal chaetotaxy;
setae used in descriptions are named; ms: s-microchaeta of thorax.
18
species and reduced from the basic pattern in Hy-
pogastruridae: sdl, sd2 and d4 are absent (fig. 7).
(6) Presence/absence of ml and a3/4 setae on th.II-
HI (thoracic segments I-II), of ml and m3 on abd.
IV and of al on abd. V. (7) Relative position of axial
setae.
The strong paurochaetosis that occurs in Odon-
tellidae results in some uncertainty about the exact
nomenclature of the remaining setae. For example,
it was difficult to say whether the second seta of the
a-row on th. II was a3 or a4; so, it was named a3/4.
Appendices and ventral chaetotaxy. — (1) Tro-
chanter. Number of setae (stable in the species
examined here). (2) Femur. Number of setae. (3)
Tibiotarsus. Number of setae (proximal + distal
whorls). Important paurochaetosis from the prim-
itive pattern 19, 19, 18 of Poduromorpha. (4) Te-
nent hair of tibiotarsus. (5) Unguiculus. Always
absent in the species studied. (6) Ventral tube. Al-
ways 3 + 3 setae in the species described here, the
same as found in the first instar of Poduromorpha
(primitive state). (7) Ventral setae of abd. I, II and
III (8) Dentes and mucro. Few noticeable variation
in our species which have always a large mucro
compared to dens. Ratio dens/mucro (d/m).
Anal spines. — Length and morphology. Anal
spines are not derived from setae, but directly from
integument secondary granules. Only S. longispina
has spine-like anal spines.
SYSTEMATIC PART
Material is deposited in the collection of Labora-
toire de Zoologie, Université Paul Sabatier (Tou-
louse).
Superodontella ciconia sp. n.
(figs. 1-7)
Type material. — Holotype young female, Doi In-
thanon, 2500 m a.s.l., humus, 2.1.81, sample n° THA 72,
Deharveng leg; 49 paratypes from numerous samples,
same station, 2000-2500 m, litter, humus and moss, 2 &
9.1.81 and 1.VIII.85, Deharveng and Gouze leg.
Description. — Colour pale blue. Length up to
1.5 mm.
Integument granulations (figs. 1 & 2). — Roun-
dish to polygonal secondary granules all over the
body, thickened on abd. VI which is globular. On
head, muscular insertions convex and 5-6 secondary
granules between d3 and d5 setae.
Antennae (figs. 3 & 4). — Ant. IV chaetotaxy as
follows: a large apical swelling of the integument;
dorso-distally 1 small ovoid organite (or) and 1 very
small s-microchaeta (ms); 9 rather long subcylin-
drical s-setae (the lateral ones slightly longer and
strongly bent), of which 7 are dorsal and 2 ventro-
lateral; dorsally and ventrally 16 long slender mou-
BEDOS & DEHARVENG: Superodontella
setae (“soies mousses”, Deharveng 1981b), 10-15
short pointed setae (including the dorsal i-seta);
ventrally 11 rather long trumpet-setae.
Ant. Ill organite composed of 2 long bent s-setae
(sl et s4), 2 swollen rods (s2 and s3) and an ex-
tremely minute, triangular, s5 microchaeta. Ant. III,
ant. II, ant. I with 15-16, 10, 6-7 ordinary setae
respectively.
Ocular area. — 5 + 5 subequal eyes; a few swel-
lings in the center of the ocular field; postantennal
organ amiboid with 4 lobes, the posterior one very
reduced.
Buccal area (figs. 5 & 6). — Buccal cone subcylin-
drical, slender and elongate. Labium with only 3
setae: F much longer than G, E very small, f absent,
without distinct apical small spines. Perilabial area
with 4 + 4 setae: b3 and b4 very long, bl and b2
shorter. Postlabial m1 seta more than twice as long
as bl; pl absent. Maxilla stipa bowed; I.P.P. and
E.P.P. apparently articulated with fulcrum arm;
LP.P. slightly longer than E.P.P.; a single mandible
present on the right side, as long as the stipa, with
arm bent at apex.
Dorsal chaetotaxy (fig. 7). — Ordinary setae sub-
equal, smooth and pointed, not long except on abd.
VI; s-setae quite similar in length and morphology
to ordinary setae, slightly thinner (position from th.
II to abd. V: 3, 3/4, 4, 4, 4, 3). On head, c2 present.
Th. II, th. IT and abd. IV with ml present, abd. V
with al present.
Appendices and ventral chaetotaxy. — As fol-
lows:
fr E 18
EI 5 10 O0
LI 5 10 6+ 10
LHI 4 9 yar Lo)
abd. I MES
abd. II Ve»
abd. III Veo)
Distal whorl of tibiotarsus with one acuminate
tenent hair. Unguiculus absent. Dentes with 5 + 5
dorsal setae, the 3 internal ones stronger; mucro
typical of the genus, as long as the dentes.
Anal spines (fig. 1). — Rather short, with strong
granular papillae.
Derivatio nominis. — The species is named for
its long and thin buccal cone.
Discussion. — Only one Superodontella species
with very long and thin mouth parts has been des-
cribed, S. cornuta Yosii, 1965 from Japan. It differs
from S. ciconta by the presence of spiniform setae
on ant. I-II and coxae, apically swollen tenent hairs
on tibiotarsi and clavate setae on abd. VI tergite.
The axial chaetotaxy of tergites is the same in the
two species but the complete dorsal chaetotaxy of
S. cornuta has not been described.
19)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
n
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D
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di
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20
Superodontella gouzei sp. n.
(figs. 8 to 14)
Type material. — Holotype male, Doi Inthanon, 2500
m a.s.l., moss on rock, 1.VIII.85, sample n° CL 47, Dehar-
veng leg; 30 paratypes from numerous samples, same
station, 2000-2500 m, litter, humus and moss, 2 & 9.1.81
and 1.VIII.85, Deharveng and Gouze leg.
Description. — Colour pale blue. Body length 0.7
to 1 mm (0.75 mm for the holotype male, 1 mm for
a paratype female, 0.7 to 1 mm for large juveniles).
Integument granulations (figs. 8 & 9). — Roun-
dish secondary granules all over the body; they are
slightly fused around setal sockets on head and on
abd. VI tergite. On head, granules ordered in two
longitudinal rows (axial area) and in oblique rows
(frontal area); 8 secondary granules between d3 and
d5; muscular insertions as fig. 8.
Antennae (figs. 10 & 11). — Ant. IV chaetotaxy
as follows: no swelling of the integument at the
apex; dorso-distally, 1 small ovoid organite (or) and
1 very small s-microchaeta (ms); dorsally and ven-
trally about 20 long mow-setae (including non dif-
ferentiated s-setae), 12-13 rather short pointed or-
dinary setae (including the dorsal i-seta); ventrally
9-10 fairly long trumpet-setae.
Ant. III organite composed of 5 ovoid short s-
setae hidden under an integument fold; sl and s4
longer than the others. Ant. III, ant. II, ant. I with
only 12-13, 10, 7 ordinary setae.
Ocular area. — 5 + 5 subequal eyes with a few
swellings in the center of ocular field; postantennal
organ amiboid with 4 lobes, the posterior one very
reduced.
Buccal area (figs. 12 & 13). — Buccal cone short
and wide. Labrum with 1 + 1 setae and a pair of
longitudinal sclerotized structures. Labium with 6
setae; G and F long, f short, E and ?A very short,
hardly distinct at higher magnification, ?d rather
short; 4 small spines on the reduced distal part.
Perilabial area with 4 + 4 setae: b2 very long, bl and
b3 subequal, b4 much smaller than the others; post-
labial ml seta slightly longer than bl; pl absent.
Maxillary stipa straight, oblique and bent at the tip
towards fulcrum; I.P.P. and E.P.P. apparently artic-
ulated with fulcrum arm; LP.P. twice longer than
Figs. 8-14. Superodontella gouzei sp. n. — 8. Central area
of head between dl and d5 setae: integument secondary
granules and muscular insertions; 9. Abdomen VI tergite;
10. Right third and fourth antennal articles, dorsal side;
i: i-seta; s5: external s-seta of ant. III organite, x: ordinary
distal seta belonging probably to ant. III; 11. Right third
and fourth antennal articles, ventral side; 12. Labral, labial
and perilabial setae, lateral view; lr: labral seta; sc: distal
sclerotized structure of labrum; sp: small distal spines; 13.
Maxilla and mandible, dorsal view; 14. Dorsal chaetotaxy;
starts: setae absent compared with pattern in S. ciconia.
BEDOS & DEHARVENG: Superodontella
Table 1. Differences between Superodontella gouzei and
S. salmoni.
S. gouzei S. salmoni
s2 and s3 setae oval, short T-shaped
on ant. III
2 anterior ocelli subequal to larger than
others others
labial setae ? A<E<f<?d<G<F ?2d,f,EG
subequal
E.P.P.; presence on the right side of a single mand-
ible shorter than the stipa, with rounded arm apex.
Dorsal chaetotaxy (fig. 14). — Ordinary setae
subequal, smooth and pointed, not long except on
abd. VI; ratio s-seta/ordinary seta decreasing from
2 to nearly 1 from thorax to abd. V (position from
th. II to abd. V: 3, 3/4, 4, 4, 4, 3). On head, c2 absent.
Th. II-III with ml and a3/4 absent; abd. IV with ml
and m3 absent, abd. V with al absent.
Appendices and ventral chaetotaxy. — As fol-
lows:
Tr E T
JE 5 11 ar)
LII 5 11 Chita?
LUI 4 10 ary)
abd. I TV:3
abd. II Ve:3
abd. III Ve:4
Distal whorl of tibiotarsus with one acuminate
tenent hair. Unguiculus absent. Dentes with 5 + 5
dorsal setae, the 3 internal ones stronger; mucro
typical of the genus, as long as the dentes.
Anal spines. — Short (fig. 9).
Derivatio nominis. — This species is dedicated to
our friend Alain Gouze who participated to the
collection of the species in 1981.
Discussion. — S. gowzet is a near relative of S.
salmoni Massoud, 1965 from New Guinea by its
reduced chaetotaxy, setae morphology, small anal
spines and short buccal cone. Differences are listed
in table 1. Type specimens of S. salmon: have been
examined but details of antennal chaetotaxy as well
as maxilla and mandible could not be observed on
this material.
Superodontella longispina sp. n.
(figs. 15-21)
Type material. — Holotype male, Doi Inthanon, 2500
m a.s.l., moss on rock, 1.VIII.85, sample n° CL47, Dehar-
veng leg; 14 paratypes from 12 samples, same station,
2000-2500 m, litter and humus, 2 & 9.1.81, 1.VIIL85 and
23.VI.86, Bedos, Deharveng and Gouze leg.
21
IES SOS
x Eri ae
ZA DITES 2 LE 2 e
Zi N In a ui n CC
ae e Neg ES \
peen ‘ SS Ci Ca !
bf mm i Pa nam, EN { L i e N = ~ A 5 em
a Ù
7 UT PA L Xe / = 4 4 &
DES sur e
ds He \ = Jan Me 74 FG } CS
n )
>
(4
sn
D
N
<
ia
BAE SN N
EEE ND =
EE 2
‘
Te
A 2 24
(
N
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
BEDOS & DEHARVENG: Superodontella
Table 2. Differences between Superodontella longispina, S. distincta, S. nepalica and S. macronychia.
S. longispina
S. distincta
S. nepalica S. macronychia
sp. n. Yosii, 1954 Yosii, 1971 Prabhoo, 1976
(Thailand) (Japan) (Nepal) (India)
apical swelling absent present present ?
on ant. IV
rows of setae 3 22 2
on abd. IV
internal teeth basal about 1/3 basal “median”
of unguis from basis
tibiotarsus long, long, acuminate, not same as
tenent hair acuminate capitate differentiated nepalica
anal spines long, long, “smaller than long, large,
pointed pointed in O. distincta”, subcylindrical
pointed and truncated
Description. — Colour grey blue, white muscular
insertion, conspicuous in alcohol. Body length up to
1.5 mm.
Integument granulations (figs. 15 & 16). —
Roundish or hexagonal secondary granules all over
the body, of small size (12 secondary granules be-
tween d3 and d5 setae on head), more convex and
thickened on abd. VI; granules joint around setae
sockets. Muscular insertions conspicuous, especially
a median longitudinal one on head as fig. 15.
Antennae (figs. 17 & 18). — Ant. IV chaetotaxy
as follows: no apical swelling; dorso-distally, 1
small ovoid organite (or) and 1 very small s-micro-
chaeta (ms); 7-8 rather long subcylindrical s-setae
(1 thinner), dorsal and ventro-lateral; dorsally and
ventrally 16 long slender 7204-setae and 15-16 short
pointed setae (including the dorsal i-seta); 31 short
ventral trumpet-setae and 2 long ones at the apex.
Ant. III organite composed of 4 T-shaped s-setae
(sl to s4) under an integument fold and an ex-
tremely minute, triangular, s5 microchaeta. Ant. IH,
ant. II, ant. I with 18, 10, 7 ordinary setae.
Ocular area. — 5 + 5 subequal eyes; about 30
secondary granules in the center of the ocular field;
postantennal organ hypogastruroid with 4 large,
well separated lobes.
Figs. 15-21. Superodontella longispina sp. n. — 15. Cen-
tral area of head between d2 and d5 setae: integument
secondary granules and muscular insertions; 16. Abdomen
VI tergite; 17. Right third and fourth antennal articles,
dorsal side; stippled setae are on ventral side of ant. III,
except sl-s4 of organite which are dorsally covered with
an integument fold; i: i-seta; s5: external s-seta of ant. III
organite; x: ordinary distal seta belonging probably to ant.
UI; 18. Right fourth antennal article, ventral side; 19.
Labial, perilabial and postlabial setae; 20. Maxilla and
mandibles, dorso-lateral view; 21. Dorsal chaetotaxy;
stars: setae absent compared with pattern in S. ciconza.
Buccal area (figs. 19 & 20). — Buccal cone
strongly protruded. Labium devoid of distal small
spines, with 4 setae: F long, the others short. Peri-
labial area with 5 + 5 setae: b3 long, bl, b2, b4 and
a fifth seta (b5) short; postlabial ml seta more than
5 times longer than bl; pl present or absent, short.
Maxilla fulcrum very strong; I.P.P. and E.P.P. appar-
ently partially articulated with fulcrum arm; E.P.P.
reduced and more or less fused to fulcrum arm; stipa
rather thin with apex bent towards the fulcrum; 2
small mandibles present with rounded apices, the
right one slightly longer.
Dorsal chaetotaxy (fig. 21). — Ordinary setae
subequal, smooth and pointed, not long, except on
abd. VI where setae are longer, stronger and acum-
inate (sometimes slightly capitated); ratio s-
seta/ordinary seta decreasing from 3 to nearly 2
from thorax to abd. V (position from th. II to abd.
V: 3, 3/4, 4, 4, 4, 3). On head, c2 present. Th. II, th.
III and abd. IV with ml present, abd. V with al
present; on th. II and th. IH, ml not aligned with
al and pl; on abd. IV, al shifted from the axis and
m3 absent.
Appendices and ventral chaetotaxy. — As fol-
lows:
ir: E ali
LI 5 12 Tats UO
LU 5 Len mar 0)
LIII 4 10 6+ 10
abd. I IVES
abd. II Mens
abd. III Ve:4
Setae of coxae nearly spiniform. Distal whorl of
tibiotarsus with one tenent hair very long and
acuminate. Unguiculus absent. Unguis with 1 basal
inner tooth and 1 + 1 strong laterobasal teeth.
Dentes with 5 + 5 dorsal setae, the 3 internal ones
much stronger; mucro typical of the genus, shorter
than the dentes (d/m = 1.6).
23
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
„ N
\ L si tel
Figs. 22-28. Superodontella flammata sp. n. — 22. Abdomen VI tergite; 23. Central area of head between dl and d5 setae:
integument secondary granules and muscular insertions; 24. Right third and fourth antennal articles, ventral side; circles:
setae present in some specimens; 25. Right third and fourth antennal articles, dorsal side; i: i-seta; s5: external s-seta
of ant. III organite (ventral); x: ordinary distal seta belonging probably to ant. III; circles: setae present in some specimens;
26. Labial, perilabial and postlabial setae; 27. Maxilla and mandible, dorsal view; 28. Dorsal chaetotaxy; stars: setae absent
compared with pattern in S. ciconia.
24
BEDOS & DEHARVENG: Superodontella
Table 3. Differences between Superodontella flammata, S. gladiolifer and S. biwonensis.
S. flammata
apical digitation of ant. IV present
thickened s-setae on ant. IV 0
c2 setae on head present
basal labial setae 3 rather long,
S. gladiolifer S. biwonensis
present absent
0 9
absent present
3 short (a fourth could ?
have been overlooked)
absent present
#3 #1,2
well developed absent
l very long
ml on th. II-III, abd. IV present
s setae/ordinary setae on th. II-III #1,2
anal spines minute
Anal spines. — Strong, with very long granular
papillae (fig. 16).
Derivatio nominis. — The species is named for
its long anal spines.
Discussion. — Three species of Superodontella,
all from Asia, have strong anal spines, large and
rather long buccal cone and T-shaped sensory rods
on ant. III like S. /ongispina. On the basis of the
original descriptions, we can give the differential
characters as mentioned in table 2. A redescription
of the species S. distincta, S. nepalica and S. macro-
nychia would be however necessary to precise their
real taxonomic status: we lack data about antennal
and dorsal chaetotaxy as well as morphology of
maxilla and mandible in these species.
Superodontella flammata sp. n.
(figs. 22-28)
Type material. — Holotype juvenile, Doi Inthanon,
2500 m a.s.l., litter and humus near a bog, 2.1.81, sample
n° THA 66, Deharveng leg; 4 paratypes in 4 samples,
same station, 2500 m, litter and humus, 2 & 9.1.81, Dehar-
veng and Gouze leg.
Description. — Colour pale blue. Body length
0.55 to 0.72 mm (no adult observed). Antennae
short and conical.
Integument granulations (figs. 22 & 23). —
Rather flat, roundish secondary granules all over
the body, enlarged on abd. VI. On head, 2 median
and 2 small lateral muscular insertions and 4 secon-
dary granules between d3 and d5 setae.
Antenna (figs. 24 & 25). — Ant. IV chaetotaxy
as follows: no apical swelling but a long distal
flame-shaped digitation; dorso-distally, 1 small
ovoid organite (or) and 1 small s-microchaeta (ms);
dorsally and ventrally 5 long slender mow-setae, 12
short pointed setae (including the dorsal i-seta); 9-
10 short ventral and 2 long apical trumpet-setae.
Ant. III organite composed of 5 short and bent s-
setae Ant Ii ant: IL, ant I with 4-16" 1026-7
ordinary setae.
Ocular area — 5 + 5 subequal eyes; postantennal
organ amiboid with 4 lobes, the posterior one very
reduced.
Buccal area (figs. 26 & 27). — Buccal cone rather
long. Labium with 4 setae: F very long, others me-
dium with G longer than E and f. Perilabial area
with 4 + 4 subequal setae; postlabial ml seta
slightly longer than bl; pl absent. Maxilla fulcrum
long and thin; distal part of LP.P. apparently artic-
ulated; E.P.P. nearly four times shorter than LPP;
LPP, fulcrum arm and stipa of equal length; pres-
ence of a single mandible on the right side, longer
than the stipa, with enlarged arm apex.
Dorsal chaetotaxy (fig. 28). — Ordinary setae
subequal, smooth and pointed, not long, longer on
abd. VI; s-setae slightly longer and thinner than
ordinary setae (position from th. II to abd. V: 3, 3/4,
4, 4, 4, 3). On head, c2 present. Th. II, th. II and abd.
IV with ml present, abd. V with al present; on th.
II and th. III, a3/4 absent.
Appendices and ventral chaetotaxy. — As fol-
lows:
IGE F T
LI 5 Jul Wars)
LII > 11 Var 2
LIII 4 10 CRIS
abd. I TV:3
abd. II Ve:3
abd. III Mes
Distal whorl of tibiotarsus with one acuminate
tenent hair, slightly differentiated. Unguiculus ab-
sent. Dentes with 5 + 5 dorsal setae, the 3 internal
ones stronger; mucro typical of the genus, nearly as
long as the dentes (d/m = 1.2).
Anal spines. — Reduced to minute integument
swellings (fig. 22).
Derivatio nominis. — The species is named for
its flame-shaped digitation at the antennal apex.
25
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Discussion. — In the presence of a long and thin
digitation at the apex of antenna, this new species
is near S. gladiolifer Massoud, 1965 from New Gui-
nea. In the other characters, it seems to be closely
related to S. biwonensis Lee, 1974 from South Ko-
rea. Differences are summarized in table 3.
REFERENCES
Deharveng, L., 1981a. La famille des Odontellidae: phy-
logénèse et taxonomie. — Travaux du Laboratoire d'E-
cobiologie des Arthropodes Edaphiques, Toulouse 3
(Me ZL,
Deharveng, L., 1981b. La chétotaxie dorsale de l'antenne
et son intérét phylogénétique chez les Collemboles
Neanuridae. - Nouvelle Revue d'Entomologie 11 (1):
3-13.
Gama, M. M. da, 1988. Filogenia das espécies de Xenylla
à escala mundial (Insecta, Collembola). - Evolucion
Biologica 2: 139-147.
Massoud, Z., 1967. Monographie des Neanuridae, Collem-
boles poduromorphes à pièces buccales modifiées. —
Biologie de l'Amérique Australe 3: 7-399.
Stach, J., 1949. The apterygotan fauna of Poland in rela-
tion to the world fauna of this group of insects: families
Neogastruridae and Brachystomellidae. - Acta Mono-
graphica Musei Historiae Naturalis, Kraków: 341 pp.
Yosii, R., 1971. Collembola of Khumbu Himal. - Khumbu
Himal 4 (1): 80-130.
Received: 21 March 1989
Revised version accepted: 6 September 1989
JEAN BELLE
Velp, The Netherlands
PROGOMPHUS NIGELLUS AND PHYLLOCYCLA
HAMATA, TWO NEW DRAGONELIES FROM BRAZIL
(ODONATA: GOMPHIDAE)
Belle, J.,
1990. Progomphus nigellus and Phyllocycla hamata, two new dragonflies from
Brazil (Odonata: Gomphidae). - Tijdschrift voor Entomologie 133: 27-30, figs. 1-10. [ISSN
0040-7496]. Published 31 July 1990.
Descriptions and illustrations are given of two new species of Gomphidae (Odonata) from
Brazil, viz. Progomphus nigellus and Phyllocycla hamata. Both male nolorypes were col-
lected in Brazil, State of Rondonia, Fazenda Rancho Grande.
Correspondence: Dr. J. Belle, Onder de Beumkes 35, 6883 HC Velp, The Netherlands.
Key words. — Brazil; Progomphus, Phyllocycla, new species.
INTRODUCTION
To the courtesy of Dr. Rosser W. Garrison at
Azusa, California, I had the pleasure of receiving for
description two new gomphid species which he had
collected himself during his investigations in the
area southwest of Ariquemes in the State of Ron-
donia, Brazil. The material in question consists of
three males; the corresponding females still remain
to be discovered. The new taxa are here published
under the names Progomphus nigellus and
Phyllocycla hamata. The holotypes are deposited in
the National Museum of Natural History, Smith-
sonian Institution, Washington, D.C; the paratype
is in the collection of Dr. Garrison.
Progomphus nigellus spec. nov.
(figs. 1-6)
Material. — Brazil: State of Rondonia, Fazenda Rancho
Grande, 62 km SW of Ariquemes (10° 50’ S, 63° 7’ W, 187
m), 2-11 November 1989, 1 @ (holotype), R. W. Garrison.
This species is the fourth member of the
guyanensis group to which also belong
Progomphus guyanensis Belle, 1966, Progomphus
approximatus Belle, 1966 and Progomphus boli-
viensis Belle, 1973. The males of this infrageneric
group differ from those of the closely allied
complicatus-group in having the superior anal ap-
pendages armed with a sharply-pointed basal ex-
terno-lateral dilatation.
Progomphus nigellus is perhaps as nearly related
to Progomphus boliviensis as to the other two
members of the group. The male superior anal
appendages are similar to those of boliviensis, the
inferior anal appendage approaches more that of
approximatus while the anal tubercles resemble
those of guyanensis. The morphological differences
and similarities in the terminalia of the abdomen
will be seen by comparing the figures of the present
species with the corresponding ones of the other
three members earlier published by me (Belle 1966,
1975);
In order not to tamper with the unique specimen
I have studied the accessory genitalia 27 situ. This
can easily be done because the posterior hamules are
in a somewhat extruded position. The accessory
genitalia then exhibit a great resemblance to those
of boliviensis.
Male (holotype). — Total length 38 mm; abdo-
men (incl. app.) 29 mm; hind wing 22.5 mm; costal
edge of pterostigma of fore wing 2.7 mm.
Smaller and more delicate than the other mem-
bers of the group. Coloration done in black, brown
and greenish white (= pale) but the pale markings
on the abdomen are yellow. Face predominantly
pale, slightly tinged brown at posterior parts of
labrum and postclypeus. Superior surface of frons
and occipital plate pale. Vertex brown. Rear of head
brown with a pair of small submedian pale spots on
top and a pair of larger pale markings on temporae.
Crest of occiput slightly and evenly concave, thinly
fringed with pale hairs which are about as long as
half the middorsal length of the occipital plate.
Prothorax with pale middle lobe and black hind
lobe. Thoracic colour pattern black with pale mark-
ings, resembling that of boliviensts but with metep-
imeral pale stripe partly developed and interrupted
in middle (fig. 1).
Legs dark brown but inner side of first femora
pale and second and third femora becoming lighter
brown toward the bases.
27
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 1-10. — 1-6, Progomphus nigellus spec. nov., male holotype: 1, diagram of thoracic colour pattern; 2, right posterior
genital hamule, ventral; 3, seventh abdominal segment, left profile; 4, tenth abdominal segment and anal appendages,
dorsal; 5, the same, ventral; 6, the same, left profile. — 7-10. Phyllocycla hamata spec. nov., male holotype: 7, diagram
of thoracic colour pattern; 8, tenth abdominal segment and anal appendages, dorsal; 9, right posterior genital hamule,
ventral; 10, apical segments of abdomen and anal appendages, left profile.
Wings with a brown tinge, blackish brown vena-
tion and brown pterostigma. Brace vein present.
Ante- en postnodal cross-veins of first series 10:14-
16:12/11-11-11:10 in fore and hind wings, respec-
tively. Second primary antenodal cross-vein the
fifth. Basal subcostal cross-vein present. All supra-
triangles one-celled. All subtriangles and triangles
two-celled with two rows of cells following but with
an extra initial cell at hind angle of triangle in hind
wings. Intermedian cross-veins 7-7/5-4 in fore and
hind wings, respectively. Anal field of fore wing
28
two cells wide for a distance of two cells. Hind
wings with five paranal cells, three (left) and four
(right) postanal cells, three rows of cells behind
Cu2, and a three-celled anal triangle.
Abdomen predominantly dark brown or blackish
brown. Terminalia of abdomen black, including
upper surface of superior anal appendages (con-
trary to the other members of the group which have
the apical part of the upper surface pale or partly
pale). Sides of segments 1 and 2 largely yellow.
Sides of segments 3 to 7 with a small, yellow basal
spot. Middorsum of segment 2 with a round yellow
spot that tapers to hind border of segment. Segments
3 to 7 with a middorsal yellow line over whole length
of segment, the line being very fine on segments 4
to 7. There is no tubercle of any sort on venter of
segment 1. Segment 7 slightly widening on apical
half of segment (fig. 3). Posterior genitalhamule and
anal appendages shaped as shown in the accompan-
ying figures. Tip of branches of inferior anal appen-
dage ending with three teeth.
In his letter of 7 April 1990, Dr. Garrison in-
formed me, that he had collected this male on a trail
at about 4:00 p.m., just before it rained, in company
with a male of Aphylla dentata Selys. In addition to
these species and Phyllocycla hamata spec. nov.
described below, he had also taken Zonophora ca-
lippus klugi Schmidt and Phyllogomphoides ce-
pheus Belle in the same environment.
Phyllocycla hamata spec. nov.
(figs. 7-10)
Material. — Brazil: State of Rondonia, Fazenda Rancho
Grande, 62 km SW of Ariquemes (10° 50 ’ S, 63° 7’ W,
187 m), 2-11 November 1989, 2 ¢ (holotype and para-
type), R. W. Garrison.
This species is the seventh member of the
volsella group (cf. Belle 1988). Dr. Garrison recog-
nized it as an undescribed species and wrote from
Azusa on 19 February 1990 to me that the males are
nearest to my armata but whose morphology of the
appendages is different. The species is peculiar by
the huge hook which arises erect on the upper
surface of each male superior anal appendage.
Male (holotype; abdomen broken between seg-
ments 4 and 5). — Total length 43 mm; abdomen
(incl. app.) 33 mm; hind wing 25.5 mm; costal edge
of pterostigma of fore wing 2.9 mm.
Head brown with pale (= leaden grey) markings.
Labrum lighter brown along free border and with
a symmetric pair of weakly developed pale spots.
External surface of genae largely pale. Anteclypeus
pale. Postclypeus with a pale spot on each lateral
side. Superior surface of frons pale, darker on the
middle line. Rear of head lighter on temporae. Crest
of occiput slightly concave, fringed with brown
hairs which are about as long as the middorsal
length of occipital plate.
Prothorax black above. Prerothorax black to dark
brown with pale (grey to grey-green) stripes; its
colour pattern shaped as shown in diagram (fig. 7).
Femora brown, but brownish yellow on inner
sides of first and second femora and on posterior
side of third femora basally. Distal spines of ante-
rior outer row of third femora more or less spaced
and one-fifth to one-fourth as long as local diameter
of femur. Tibiae, tarsi and claws black.
BELLE: New dragonflies from Brazil
Wings with a brown tinge, black venation and
light brown pterostigma. Brace vein present. Ante-
and postnodal cross-veins of first series 12:17-
15:13/11:12-12:11 in fore and hind wings, respec-
tively. Second primary antenodal cross-vein the
fifth. Basal subcostal cross-vein present in all wings.
Supratriangle in right hind wing three-celled, in
other wings two-celled. Subtriangles in fore wings
two-celled, in hind wings one-celled. Triangles two-
celled with two rows of cells following but with an
extra initial cell at hind angle of triangle in hind
wings. Intermedian cross-veins 9-8/6-6 in fore and
hind wings, respectively. Anal field of fore wings
two cells wide for a distance of two cells in anterior
row. Hind wings with four paranal cells and three
postanal cells (the fourth paranal cell is the first
postanal cell), a one-celled anal loop, three rows of
cells behind Cu2, and a four-celled anal triangle.
Abdomen dark brown with paler markings as
follows: Sides of segments 1 and 2 largely yellow.
Sides of segments 3 to 7 with yellow baso-lateral
spots, the basal spots of both sides of segment 7
connected on the middorsum at extreme base.
There is a yellow middorsal line on segments 2 to
7. Sides of segments 8, 9 and 10 largely brown-
yellow. Lateral dilatations of segments 8 and 9 nar-
row and black. Lateral margins of segment 8 with
four (left) and three (right) denticles at apex, those
of segment 9 denticulated at extreme base. Poste-
rior margin of segment 10 denticulated at level of
bases of superior anal appendages. Dorso-apical
rim of segment 10 about one-fourth the middorsal
length of segment. Superior anal appendages black
and shaped as shown in figs. 8 and 10. Rear margin
of vesicle deeply cleft, bottom of cleft rounded and
without a median elevation of any sort. Tip of
posterior genital hamule rather long (fig. 9).
The male paratype lacks the tip of the left super-
ior anal appendage but for the rest the specimen is
in a perfect condition. The pale markings of the
pterothorax are yellowish green and the dark lateral
stripes brown. The wings are slightly brown-
tinged. The cubito-anal interspace of the right hind
wing has two cross-veins instead of the usual single
cross-vein and the pterostigma is a trifle larger than
that of the male holotype. The measurements of the
male paratype are: Total length 42.5 mm; abdomen
(incl. app.) 32.5 mm; hind wing 24.5 mm; costal
edge of pterostigma of fore wing 3 mm.
Remark. — I take this opportunity of correcting
a misprint in the description of Phyllocycla armata
Belle, 1977. On page 7, in rule 7 from above, part
of a line has been left out. We have to read: “Trig-
onal interspace in fore wings starting with two
rows of cells from triangle outwards, that in hind
wings starting with a row of three cells against
triangle followed by two rows of cells.”
29
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
REFERENCES
Belle, J., 1966. Surinam dragon-flies of the genus
Progomphus. - Studies on the fauna of Surinam and
other Guyanas 8: 1-28; pls. 1-4.
Belle, J., 1973. A revision of the New World genus
Progomphus Selys, 1854. - Odonatologica 2: 191-308.
Belle, J., 1977. A new species of Phyllocycla Calvert, 1948
from Brazil (Odonata: Gomphidae). - Entomologische
Berichten, Amsterdam 37: 6-7.
30
Belle, J., 1988. A synopsis of the species of Phyllocycla
Calvert, with descriptions of four new taxa and a key
to the genera of Neotropical Gomphidae (Odonata,
Gomphidae). - Tijdschrift voor Entomologie 131: 73-
102.
Received: 17 May 1990
Accepted: 17 May 1990
K. A. MONK & R. K. BUTLIN*
Department of Pure and Applied Zoology, University of Reading.
* School of Pure and Applied Biology, University of Wales.
A BIOGEOGRAPHIC ACCOUNT OF THE
GRASSHOPPERS (ORTHOPTERA: ACRIDOIDEA) OF
SULAWESI, INDONESIA
Monk, K. A. & R. K. Butlin, 1990. A biogeographic account of the grasshoppers (Orthoptera:
Acridoidea) of Sulawesi, Indonesia. — Tijdschrift voor Entomologie 133: 31-38, tabs. 1-4.
[ISSN 0040-7496]. Published 31 July 1990.
Nineteen acridid species were collected from the Dumoga Bone National Park and its
environs in North Sulawesi, Indonesia, during 1985: two were new to science and one was
known previously only from the Philippines.
The total acridoid fauna of Sulawesi, including these three species, comprises 61 species of
Acrididae (53 Catantopinae, four Oedipodinae, three Acridinae, one Gomphocerinae) and
one species of Pyrgomorphidae. This fauna has four main characteristics: (a) a high level
of specific and generic-level endemism, (b) low generic-level diversity but a high number
of species per genus, (c) diverse distributions outside Sulawesi, and (d) localised distributions
within the island.
In the Catantopinae, 83% of species are endemic to Sulawesi, reflecting the importance of
the island as an area of endemism. Species distributions within Sulawesi suggest that each
region of the island (North, Central, South and South-east) may also be an area of endemism
in its own right.
These observations are discussed in relation to the geological history of Sulawesi. They
suggest that for much of its history the island has been more isolated from other land masses
than it is at present, and that it has been fragmented either into separate islands or by
ecological barriers.
Correspondence: Dr. R. K. Butlin, School of Pure and Applied Biology, University of Wales,
P.O. Box 915, Cardiff CF1 3TL, Wales.
Key words. - Biogeography, Sulawesi, Wallacea, grasshoppers, endemism.
INTRODUCTION
Although the Acridoidea in temperate zones and
in dry equatorial areas are well-known, only wide
ranging, economically-important pests, e.g. Oxya
japonica japonica Thunberg and Locusta migratoria
Linnaeus, have been studied in wet equatorial areas.
The taxonomic work in South-east Asia by Ramme
(1941) and Willemse (1951, 1956, 1957) is an ex-
ception but it has not been extended. In particular,
the acridoid fauna of Sulawesi has been largely ig-
nored despite the great interest in other animal
groups inhabiting the island.
Sulawesi lies in a deep sea region, also including
the Lesser Sunda Islands (Lombok to Timor) and
the Moluccas, with lies between two continental
margins: the Sunda (Malaysia and the Greater
Sunda Islands), and the Sahul shelves (New Guinea
and offshore islands). Classically, this area was seen
as the meeting point of two major biotas: the Orien-
tal and the Australasian. Wallace (1869) originally
separated them by dividing Bali (and Borneo) from
Lombok (and Sulawesi). Although Wallace’s Line
may apply to some groups such as freshwater fish
and mammals, the diversity of biogeographical
patterns across this region for different taxonomic
groups is high (see George 1987). Even with this
overlap of broad transitional zones, Wallacea (a
collective term for the islands of the deep sea re-
gion) is generally considered to be insular and bi-
otically impoverished, but with unique characteris-
tics indicating long periods of isolation (Dickerson
et al. 1928).
Sulawesi covers 159,000 km? divided into four
regions, Utara, Tengah, Selatan, and Tenggara, or
North, Central, South, and South-east Sulawesi, re-
spectively (see Whitten, Mustafa & Henderson
1987, for geographical details). North Sulawesi ts a
narrow peninsula, 772 by 103 km. Volcanoes, 1800-
2400 m above sea level (asl), range along the penin-
sula. Seismic activity was recorded during 1985 and
31
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
1986. Limestone outcrops (possibly Cretaceous) oc-
cur in the western half of North Sulawesi. The
northern peninsula is linked with Central Sulawesi
by a narrow neck of land at the western extremity.
The Sangihe and Talaud Islands lie between the
northern tip of North Sulawesi and the Philippines.
Central Sulawesi is also mountainous, with peaks
reaching 2000-3000 m asl. A limestone mountain
range running north to south through South Su-
lawesi is 500-1000 m asl and bisected by a narrow
plain. The lakes on the plain are only 1-2 m deep
and were once part of the sea, suggesting that at
least this peninsula may have been partially sub-
merged at times. The single volcano in this region,
Lompobatang (2871 m asl), is extinct. The penin-
sula of South-east Sulawesi has a western mountain
range reaching 2800 m asl and a lower-lying eastern
area.
The vegetation is affected by regional climatic
differences. Central Sulawesi has the greatest ratio
of wet to dry months, whereas more seasonal rain-
fall occurs in the peninsulas (Whitmore 1984). The
central region is covered in evergreen rain forest,
whereas semideciduous forest is more common in
the peninsulas (Walker 1982). Open habitats are
mainly secondary, due to forest clearance for agri-
culture (Whitten et al. 1987).
Geologically Wallacea is a highly complex region
originating in an interaction between the South-
east Asian, Australian, and Pacific plates (Charlton
1986). The present island of Sulawesi is believed to
be derived from two fragments that originated se-
parately during the breakup of east Gondwanaland
about 320 My ago, and remained separate during a
northerly move starting 220 My ago (Audley-
Charles 1987). From the late Cretaceous onwards,
these fragments probably formed part of an ar-
chipelago between the Asian mainland and Austra-
lia-New Guinea but their positions relative to other
land masses are uncertain. Holloway (1987) em-
phasized two alternative theories about the rela-
tionship of Sulawesi with Borneo: 1. The fragment
now forming the western part of the island (North,
South and part of Central Sulawesi) has always been
in about its present position relative to Borneo,
whereas the eastern fragment was Australasian in
origin, 2. The two fragments were both isolated
island arcs, now fused and thrusted towards Borneo.
The present island was probably formed about 15
My ago. The area of land above sea level, its division
into separate islands and its climate are additional
factors which must have contributed to the present
day fauna of Sulawesi but about which little is
known (Morley & Flenley 1987).
It is now believed that Sulawesi was never linked
by land bridges to any other land mass following its
formation 15 My ago (Audley-Charles 1987; Hollo-
way 1987). During the Pleistocene, the sea level fell
several times when affected by the Glacial expan-
32
sion of the ice caps. The largest drop, 170,000 years
ago, linked Sundaland (Sumatra, Java, Bali, and Bor-
neo) with the Asian mainland, with land extensions
northward to the Philippines. Exposed land also
extended around South Sulawesi toward the Lesser
Sunda Islands, and from Central Sulawesi eastward
toward New Guinea. During these periods of low
sea level a drier, more seasonal climate prevailed
and dry savannah may have extended through the
Philippines to the Lesser Sunda Islands and into
Australia, separating two large, wet rainforest areas
in Sundaland and New Guinea; some seasonal
plants still survive in South Sulawesi (van Steenis
1979). The climatic fluctuations and corresponding
vegetational changes of the Quaternary are now
known to have occurred in the late Tertiary as well
(Morley & Flenley 1987).
MATERIALS AND METHODS
The analysis reported here is based on a combi-
nation of data from the literature and collections
made by the authors and others in the Dumoga-
Bone National Park, North Sulawesi during 1985.
This work formed part of the Royal Entomological
Society's ‘Project Wallace’ expedition. Insects were
collected by sweeping, beating, and hand-searching.
Agricultural crops, grassland, and primary semi-
deciduous forest (undergrowth, bushes, saplings,
and trees up to 2 m) were examined at all times of
day and night and in three separate periods cover-
ing most of the year. Specimens collected from the
tree canopy during the British Museum (Natural
History) fogging programme (N. Stork, unpub-
lished) were also examined.
The principal published records of the acridid
fauna of Sulawesi are: Ramme (1941), Dirsh
(1954), Willemse (1951, 1956, 1957, 1968), Hollis
(1968, 1971, 1975), Kevan & Chen (1969), and
Ritchie (1982). Family and subfamily classification
follows that adopted in the collection of the British
Museum (Natural History).
RESULTS
Expedition collection of North Sulawesi
Acridoidea
Nineteen species from eighteen genera of Acri-
doidea were found in the Dumoga Bone National
Park and its environs during 1985 (table 1). This
includes two undescribed catantopine species, from
the genera Tarbaleus and Bibracte, both of which
were collected from the canopy by fogging. One
species, Eoscyllina luzonica, was previously known
only from the Philippines and was collected only
from limestone grassland in the western part of the
National Park. This suggests that the data available
in the literature on the species present in Sulawesi
MONK & BUTLIN: Grasshoppers of Sulawesi
Table 1. Species of Acridoidea occurring in Sulawesi and their distributions.
Wings! Distribution?
Outside Within
Pyrgomorphidae
Atractomorpha Across
psittacina psittacina de Haan ah W SYP All
Acrididae
Acridinae
Acrida Across
willemsei Dirsh + W SWP All
Calliphlaeoba Sulawesi
celebensis Ramme ar B — N,C
Phlaeobacris Sulawesi
reticulata Willemse W — €
Gomphocerinae
Eoscyllina P
luzonica Bolivar I SF W P N È
Oedipodinae
Heteropternis Across
obscurella Blanchard W Across All
Atolopus Across
thalassinus tamulus Fabricius Ar W Across All
Locusta Across
migratoria Linnaeus ali W Across All
Gastrimargus Across
marmoratus (Thunberg) W Across All
Catantopinae - Endemic genera
Alectorolophus deceptor Ramme ala B _ SE, N >
obscoenus Brunner von Wattenwyl U B — N =
unilobatus Brunner von Wattenwyl B — S 5
applicatus Brunner von Wattenwyl B — ISTE È
speciosus Brunner von Wattenwyl B — S
mutator Ramme B — N È
sororum Ramme B _ E =
lineatus Ramme B — C Bs
guttulosus Ramme B = N 5
Acrolophus cornutus Ramme B — C ES
Alectorolophellus heinrichi Ramme B — S 5
Mengkokacris olivacea Ramme W? _ SE =
Heinrichius nobilis Ramme W — N >
Celebesia acuticerca Bolivar C ala W? — N x
ferruginata Brunner von Wattenwyl We? — N Ri
heinrichi Ramme W? = S à
Paramesambria flavomaculata Willemse B — S
Paracranae celebesia Willemse Ar B — N ©
Catantopinae - Non-endemic genera
Tarbaleus Brunner von Wattenwyl M, NG
sp. n. te B — N =
Oxya Across
Japonica japonica Thunberg W SP All
bolaangensis Hollis W -- N “
stresemanni Ramme W — C sa
Gesonula Across
mundata pulchra Rehn SF W S N
Chitaura M, Java, India?
brachyptera Bolivar I B N
flavolineata (Willemse) B — N Ea
atrata Ramme B — C 4
mirabilis Carl B C 5
ochracea Ramme B _ S Sp
vidua Carl B M S
mengkoka Ramme B — SE
samanga Carl B _ SYSEAE
poecila Ramme B — N
elegans Ramme B — S +
Ue
(US)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 1. Species of Acridoidea occurring in Sulawesi and their distributions. (continued).
Oxytauchira
gracilis Willemse
Stenocatantops
splendens Thunberg
angustifrons Walker
Bibracte Stàl
sp. n.
Valanga
transiens Walker
Oxyrrbepes
meyeri Willemse
obtusa de Haan
Cranaella
carnipes Ramme
Tristria
pisciforme Serville
Austracris
guttulosa guttulosa Walker
Mesambria
maculipes Stàl
elegans Ramme
trapezina Ramme
rectangularis Ramme
Traulia
kukenthali Ramme
sanguinipes Stàl
Apalacris
gracilis Willemse
incompleta Willemse
celebensis Willemse
Moessonia
tenebrifera Walker
Notes
+ indicates a species collected by Project Wallace
1 - W - fully winged
W? — winged but probably a poor flier
B - brachypterous or wingless
2 - Distributions outside Sulawesi:
M - Moluccas
S — Sundaland
P - Philippines
A - Australasia
NG - New Guinea
Across - at least Sundaland to New Guinea
are reasonably reliable since intensive collecting
produced so few undescribed species, and those only
from inaccessible habitats. Data on distribution
within Sulawesi is less reliable with a significant
proportion of species described from only one or a
few localities. However collecting localities in the
past have been widespread and so this may reflect
very localised species distributions (see below). A
probable exception is Central Sulawesi, parts of
which remain very inaccessible. The expedition col-
lection represents only a small proportion of the
total acridoid fauna of Sulawesi (16 out of 59
described species) and this is also likely to be a result
34
Wings! Distribution?
Outside Within
Burma
W = C *
Across
ala W Across all
W Across S
S, M, P
+ B = N =
Across
Ww = N,C
S
W = ? *
W S N, C
P
B — N È
S
W S N
PA
W A N
India?
+ B — all
B S
B = @ =
B = N +
S,P, M
Ww? pus > *
We? S ?
S,P
W = S
W = S
Ww wee ? *
NG
We = N
2 - Distributions within Sulawesi:
all - all four regions
N - North
S - South
SE - South-East
€ - Central
* indicates that the species is known only from one
or a few sites.
of localised species distributions rather than under-
collecting.
The acridoid fauna of Sulawesi
Expedition and literature data give a total of 62
Sulawesi acridoid species in 33 genera and five sub-
families (table 1). The Catantopinae form by far the
largest group with 53 species. This very diverse
subfamily, which some authors consider to be an
amalgam of several distinct subfamilies (eg Dirsh
1961), is one of the few acridoid groups to have
adapted to forest environments. All of the non-
catantopine species in Sulawesi are forest edge or
open habitat species whereas about 80% of catan-
topine species occupy habitats within the forest.
Related to this is the occurrence of flightless spe-
cies, with wings either absent or reduced, which are
much more common in the Catantopinae (37/53
species) than in the other subfamilies (1/9 species).
Flightlessness in grasshoppers is probably an adap-
tation to poor habitat quality, diversion of resources
away from wing and flight muscle production al-
lowing earlier reproduction and/or greater fecun-
dity (Ritchie et al. 1987), and is common in forest
species in other areas (Jago 1973).
Flightlessness clearly limits dispersal and so may
have an impact on levels of endemism. In Sulawesi
endemism is high for the Acridoidea, as it is for
many other groups (Whitten et al. 1987) including
Fumastacidae and Tetrigidae in the Orthoptera
(Butlin et al 1989, Blackith & Blackith 1988). Over-
all 74% of acridoid species are endemic to Sulawesi
but the proportion is higher in the Catantopinae
(83%) than in the other subfamilies (22%). There
is a clear link with the ability to fly: only two out
of nine non-endemic catantopines are flightless,
and the one flightless non-catantopine is endemic
(table 1).
Table 2. Comparison of the Catantopinae of Sulawesi and
Java.
Java Sulawesi
Number of species recorded 47 DI,
Number of genera recorded 34 24
Number of species per genus 1.58 DAI
Proportion of endemics — species 40% 83%
- genera 15% 33%
Table 3. Distributions outside Sulawesi.
Non- Non- Relationships
endemic endemic of Endemic
Species! Genera? Genera’
Across >) 1 (10) 0
Sunda Shelf 7 4 (8) À
Borneo 0 0 (0) 1
(exclusively)
Philippines | 1(1) 0
(exclusively)
Moluccas I 0 (0) 0
(exclusively)
Australasia l 2 (3) 1
Notes
1. Includes distributions of Oxya japonaci japonaci, Ge-
sonula mundata pulchra, and Austracris guttulosa gut-
tulosa. All species have distributions ‘Across’ but the
subspecies have informative distributions.
2. Excluding (including) genera with non-endemic spe-
cies.
3. See text for details.
MONK & BUTLIN: Grasshoppers of Sulawesi
The special nature of the Sulawesi fauna can be
appreciated better if it is compared with the fauna
of Java. Java is the most comparable island in the
region in terms of land area (126,500 km?) but is
part of the Sunda Shelf and has a clearly Oriental
fauna. This comparison can be made most easily for
the Catantopinae using data from C. Willemse
(1956, 1957), E Willemse (1965) and Hollis (1971,
1975). The total number of species recorded is sim-
ilar (table 2) but the number of genera present in
Sulawesi is lower with a significantly greater
number of species per genus (X? = 6.5 P < 0.01)
Endemism at the species level in Sulawesi is twice
that in Java, as is generic level endemism.
Biogeographical links
Sharing of taxa between Sulawesi and its sur-
rounding areas is summarised in table 3 for non-
endemic species (and subspecies) and genera. It is
clear that Sulawesi shares the greatest number of
species and genera with the Sunda Shelf as a whole
(Peninsula Malaysia, Borneo, Sumatra, Java and
Bali). This bias is strongest at the specific level but
is also present among non-endemic genera. We
have found no exclusive links with Borneo at either
level, despite the close proximity of the two islands.
Links with the Philippines are mostly due to
species or genera which also occur on the Sunda
Shelf but there are two exclusive links, the gom-
phocerine Eoscyllina luzonica and the catantopine
genus Cranaella. No species shared between Su-
lawesi and the Lesser Sunda Islands have been iden-
tified. To the east there is one species, Chitaura
vidua, which occurs in Sulawesi and the Moluccas
and one, Austracris guttulosa, which is distributed
widely in Australasia and for which Sulawesi is the
western limit. At the generic level there are two
further links to the east, the genera Moessonia and
Tarbaleus.
For endemic genera biogeographical information
can only be derived from the distribution of sister
genera but since the phylogeny of these groups has
been studied very little these relationships are not
known with any certainty. The following relation-
ships, suggested by Ramme (1941) and Willemse
(1951, 1956, 1957), are included in table 3. A group
of four endemic genera, the Acrolophi (Acrolophus,
Alectorolophus, Alectorolophellus and Mengkok-
acris), has apparently evolved within Sulawesi, in-
dicating a long period of isolation. The nearest
relatives of this generic group are apparently
Paralectorolophus and Lyrolophus which occur on
Lombok and Java respectively. The acridine genera
Calliphlaeoba and Phlaeobacris are both probably
related to Phlaeoba which has a Sunda Shelf plus
Philippines distribution.
Heinrichius is believed to be most closely related
to Noliba and therefore provides the only exclusive
35
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
link with Borneo. Paracranae is part of the Cranae
group of genera which occurs in the Moluccas and
New Guinea (Willemse 1977a, b).
Two genera are problematic because their near-
est relatives are non-endemic Sulawesi genera. This
implies that the latter are polyphyletic since the
Sulawesi representatives of the two genera of each
pair are likely to be more closely related to one
another than the Sulawesi species of the non-en-
demic genus are to species in the same genus occur-
ing elsewhere. Celebesia is believed to be most
closely related to Traulia (Sulawesi and the Sunda
Shelf), an Paramesambria to Mesambria (Sulawesi
and India, Henry 1942). The distribution of
Mesambria is particularly surprising but it is paral-
leled by Chitaura which also has an Indian represen-
tative (Hollis 1975). Neither of the two endemic
genera (Celebesia and Paramesambria) has been
included in table 3.
Endemism within Sulawesi
Mosaic distributions of species or subspecies
within Sulawesi have been noted for several animal
groups, eg macaques, carpenter bees, pond skaters
(Whitten et al 1987 and see Knight & Holloway
1990). Ramme (1941) commented on a similar
phenomenon in the Catantopinae and this appears
quite striking when the island is divided into re-
gions (table 4). A high proportion of the species
present in any one region are known only from that
region. This is particularly clear in the two most
species rich genera in Sulawesi: Alectorolophus has
nine species, seven of which are known from only
one region, and Chitaura has ten species, nine of
which are known from only one region. However
the distributional information available is limited.
Many species (table 1) are known from only one
locality, but collecting localities overall have been
widely distributed around the island and so this may
represent genuinely very restricted distributions.
Where more localities are known, five out of seven
endemic and six out of eight non-endemic species
are restricted to one region. Intensive searching of
a restricted area during the Project Wallace expedi-
tion yielded only a small proportion of the total
number of catantopine species known to occur in
Table 4. Regional distribution of Catantopine species
within Sulawesi.
Region Number of species occurring in:
that region only that region and
elsewhere
North 19 5
Central 8 6
South 12 5
South-East 3 5
36
Sulawesi (10/53) but a much higher proportion of
the species known to occur in North Sulawesi
(9/21). Only one species previously reported from
another region was collected. Nine of the species
not collected on the expedition were previously
reported only from the western part of the penin-
sula of North Sulawesi, (particularly Toli-Toli) dis-
tant from the Dumoga-Bone National Park, sug-
gesting that distributions are also restricted within
regions. The within island endemism in the genus
Chitaura is particularly striking in this context. Spe-
cies in this genus are brightly coloured and conspic-
uous and occur on the forest edges as well as in light
gaps within the forest. It is unlikely that the expe-
dition would have failed to collect any species occur-
ring in the Dumoga Bone area and yet only one of
the ten Sulawesi species was found. This argument
applies to collecting of Chitaura ın general and so
this genus gives strong support to the existence of
intra-island endemism, and would be a good candi-
date for further work.
DISCUSSION
The Acrididae of Sulawesi show four main char-
acteristics: (a) a high level of endemism, (b) low
generic diversity but a high number of species per
genus, (c) diverse distributions outside Sulawesi,
and (d) localised species distributions within the
island. These features have parallels in several
other animal groups (Whitten et al 1987, Knight &
Holloway 1989) and can be related to the geological
history of the island.
The Acridoidea did not begin their radiation until
the mid-Tertiary (Sharov 1968), that is after the
break up of Gondwanaland and the beginning of the
proposed northward movements of the fragments
of present day Sulawesi. Thus the current Acridoid
fauna of Sulawesi must have originated by dispersal
and its low generic diversity can be explained in one
of two ways: either Sulawesi was more isolated for
much of its history than it is at present, or it has
only recently become dry land due to tectonic uplift.
The former explanation is more consistent with the
observation of high levels of specific and generic
endemism and high numbers of species per genus
on the island, all of which suggest a long period of
independent evolution for the Sulawesi acridoids.
The sources of colonists for Sulawesi appear to have
been numerous although the largest number of
shared species and genera is clearly with the Sunda
Shelf region. This may be partly due to the higher
diversity of Acridoidea in Asia than in Australasia
but there are several species or subspecies distribu-
tions which extend eastwards only as far as Sulawesi
and very few equivalent Australasian distributions
which extend westwards to Sulawesi. Thus at least
a part of Sulawesi appears to have been closer to the
Sunda Shelf than to Australasia for a significant
part of its history. However its current proximity
to Borneo is probably recent since there are no
exclusive biogeographical links at the level of
shared species or genera. This is also true in other
animal groups, notably the butterflies (Vane-
Wright 1990). Exclusive links with the Philippines
and Moluccas are also weak and may be recent. This
is consistent with a general geological view of the
area as a collision zone in which the area of dry land
is increasing and land masses are moving closer
together.
The high proportion of endemic species and
genera in Sulawesi, the number of species per ge-
nus, and the evidence for evolution of new genera
within the island all suggest not only a long period
of isolation but also the opportunity for consider-
able evolutionary change and speciation. The mos-
aic distributions of species, particularly in the forest
dwelling, flightless catantopines, may provide a
clue to the conditions which have favoured these
developments. Mosaic distributions are a feature of
several other animal groups (Whitten et al 1987,
Knight & Holloway 1990) and the patterns in dif-
ferent groups appear to be broadly coincident, de-
fining areas of endemism within Sulawesi. The
simplest explanation for these areas of endemism
is past fragmentation of Sulawesi. This could have
been in the form of an archipelago at times of
higher sea level or before tectonic uplift and vol-
canic activity had created the present land mass
(Musser 1987), or due to reduction in the extent of
forest habitats in periods of drier climate (Morley
& Flenley 1987). The wingless, forest adapted ca-
tantopines would be particularly susceptible to pop-
ulation fragmentation by such processes. Evolu-
tionary divergence on islands or in refugia would
have been followed by expansion to form the pres-
ent mosaic distribution. Contact zones between the
forms have not been studied, or indeed accurately
located, and so the consequences of renewed contact
are unknown. In general several possible types of
interaction exist: sufficiently divergent forms may
be able to spread into sympatry progressively obs-
curing the mosaic pattern, less divergent forms may
interact at sharp parapatric boundaries due to eco-
logical exclusion, or where reproductive isolation is
incomplete a hybrid zone may form (Barton & He-
witt 1985). Identification and analysis of such con-
tact zones is likely to be a very productive area for
future research.
Strict isolation on islands or in refugia may not
be necessary for evolutionary divergence (Barton
1989), especially in an island with the curious shape
of Sulawesi in which gene flow between popula-
tions inhabiting different peninsulas would be ex-
tremely restricted even if they were connected by
continuous suitable habitat. However,divergence
without isolation is likely to produce different pat-
terns of variation for individual characters within
MONK & BUTLIN: Grasshoppers of Sulawesi
species as well as for different species. Coincidences
of patterns of variation are most likely to be pro-
duced by contraction and expansion of populations,
and this process may be necessary for the comple-
tion of speciation (Hewitt 1989).
The mosaic distributions of species within Su-
lawesi constitute evidence either for the archipe-
lagic nature of the land mass in the past or for
habitat fragmentation during climatic fluctuations,
or both. The time scale for these changes, and thus
for the evolution of the Sulawesi endemics, is un-
certain. They could have occurred mainly in the
Pleistocene or over a much longer time scale of the
order of the 15Myrs suggested as the time since the
formation of Sulawesi as a single unit (Audley-
Charles 1987). Most probably the present fauna of
Sulawesi is the result of several superimposed pe-
riods of divergence. Studies of genetic divergence
and detailed investigations of the contact zones
appear to be the way forward in understanding the
evolution of the Sulawesi grasshoppers.
ACKNOWLEDGEMENTS
We thank Judith Marshall (BM(NH)) for much
help in the identification of specimens, and Drs J.
Holloway, R. Vane-Wright, H. Malicky, D. Clayton,
and M. Robertson for comments on earlier drafts of
the manuscript. We are grateful to Prof. R. Blackith
for the use of his unpublished data. KAM thanks
the Royal Society, the Godman Exploration Fund,
and the Percy Sladen Memorial Fund for financial
support. RB thanks the Percy Sladen Memorial
Fund, Mr C. R. Butlin and the School of Biological
Sciences, University of East Anglia.
This paper is based on material collected whilst
the authors were participants on Project Wallace,
sponsored by the Royal Entomological Society of
London and the Indonesian Institute of Sciences
(Results of Project Wallace No. 45).
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Received: 7 July 1989
Accepted: 19 September 1989
THOMAS PAPE
Zoologisk Museum, Copenhagen
TWO NEW SPECIES OF SARCOPHAGA MEIGEN
FROM MADEIRA AND MAINLAND PORTUGAL
(DIPTERA: SARCOPHAGIDAE)
Pape, T., 1990. Two new species of Sarcophaga Meigen from Madeira and mainland Portugal
(Diptera: Sarcophagidae). — Tijdschrift voor Entomologie 133: 39-42, figs. 1-9. [ISSN
0040-7496]. Published 31 July 1990.
Sarcophaga (Discachaeta) amputata sp.n. (Madeira) and Sarcophaga (Pierretia) iulicida
sp.n. (Portugal) are described and their phylogenetic affinities are discussed. S. zwlicida was
bred from the millipede Ommatoiulus moreleti (Lucas, 1860).
Dr. T. Pape, present address: Danish Bilharziasis Laboratory, Jaegersborg Allé 1D, DK-2920
Charluttenlund, Denmark.
Key words. - Diptera; Sarcophagidae; New species; Madeira; Portugal.
INTRODUCTION
A number of specimens representing two undes-
cribed species of Sarcophaga Meigen were reco-
vered from various sources. As the accompanying
biogeographical and biological information is
rather interesting, it is felt that the following des-
criptions are justified, although they do not form
part of a more thorough revision.
Depositories for specimens examined are indi-
cated by the following acronyms:
BMNH — British Museum (Natural History), Lon-
don, England; NMwc - National Museum of Wales,
Cardiff, England; ZMUC - Zoologisk Museum, Uni-
versity of Copenhagen, Denmark.
SYSTEMATIC PART
Sarcophaga (Discachaeta) amputata sp.n. (figs. 1-4)
Type material. — Holotype @, Madeira (Portugal): Be-
tween Canical & Prainha, 5.viii.1981, M. Jones (NMWC).
The holotype is in good condition, glued to the pin along
the right side of thorax and with the terminalia dissected
and glued to a piece of cardboard pinned with the specimen.
Paratypes. Madeira: 18 19, [no further locality, but
with the number 1948-301], T.W. Wollaston (BMNH,
NMWC); 29, data as holotype (NMWC, ZMUC); North
end of Deserta Grande, 14 19, 18.viii.1981, M. Jones
(ZMUC, NMWC).
Description
Male. — Head. Narrowest part of frons
0.25-0.30 X head width. Parafacial plate with a
uniserial row of setae along the eyemargin, the
lower setae of this row being long and bristly. Post-
gena with white setae in about posterior 0.50-0.75
(strict lateral view).
Thorax. Proanepisternum setose in about upper
half. Chaetotaxy: 3 indistinct pairs of presutural
acrostichals, postsutural acrostichals absent or re-
presented by the prescutellar pair only, dorsocen-
trals = 3 + 3, intra-alars = 1 + 2, supra-alars = 1
+ 3, post alars = 2. Scutellum with a pair of apicals,
3-4 pairs of marginals and without discals. Mid
femur with 2-3 anteroventral bristles, no postero-
ventral bristles. Hind tibia with a sparse row of
elongated posteroventral setae.
Abdomen. Tergites 3-4 with silvery grey micro-
tomentum mainly restricted to anterior half and
almost non-changing; interrupted by a median
black stripe. Tergite 5 with microtomentum re-
duced to narrow strips in antero-lateral position.
Terminalia black or blackish brown. Protandrial
segment with a row of marginal bristles. Cercus in
profile with a distinct subapical dorsal hump. Ae-
deagus with very long harpes that terminates in a
distinct hook, and a long, terminally bifid juxta.
Gonopod with the usual row of long setae along
dorsal margin but with some additional setae on the
median surface.
Length: 8-9 mm (estimated from dissected spe-
cimens).
Female. — Very like the male apart from the
usual sexual dimorphism. Abdominal pattern
slightly more changing and with the median black
stripe almost absent. Mid femoral organ not differ-
entiated. Terminalia red. Tergite 6 broadly arched,
the dorsal part reduced to a narrow strip, and mar-
ginal bristles only present laterally.
Length: 7.0-7.5 mm.
39
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Distribution. — Palaearctic: Madeira islands
(Madeira, Deserta Grande).
Etymology. — A Latin adjective, amputatus = cut
off, amputated. The name refers to the structure of
the aedeagus where the pair of very short arms at
juxtal base will separate the species from all other
species of the subgenus Discachaeta.
Remarks. — The sarcophagid fauna of Madeira
was reviewed by Pape (1986), who listed a total of
five species, three of which were judged to be recent
introductions by man. The present species is inter-
esting as it is the third apparently endemic species,
and much evidence is in favour of considering this
the sister species of the Madeiran Sarcophaga
(Discachaeta) kunonis (Pape, 1986). The latter spe-
cies was described in the genus Discachaeta Ender-
lein, which here is given subgeneric rank following
the wide concept of Sarcophaga employed by Séguy
(1941), Downes (1965) and Pape (1988). Disca-
chaeta is probably a monophyletic group, this hy-
pothesis being corroborated by at least the follow-
ing character states that all seem to be derived with
CX h
Figs. 1-4. Sarcophaga amputata, male
terminalia. 1, right cercus and surstylus,
lateral view, setae omitted; 2, cerci, pos-
terior view, setae omitted; 3, right par-
amere + gonopod, lateral view; 4, ae-
deagus, lateral view. Abbreviations: h
= harpes, j = juxta, p = process at
juxtal base. Scale = 0.1 mm.
40
regard to the groundplan of the Sarcophaginae and
probably with regard to that of Sarcophaga sensu
lato as well: 1) Male hind coxa with a ventro-
median pad of short spiny bristles; 2) Aedeagal
juxta more or less dome-shaped and arching over
the lateral styli; 3) Base of juxta with a pair of arms
or processes; 4) Male cercus dorsally with a flat-
tened or concave area and with a more or less
distinct hump subapically; 5) Female mid femoral
organ, if present, in apical position.
Defined in this way, Discachaeta contains six
species, all distributed in the western Palaearctic.
(Note that the species Discachaeta gigas Povolny,
1986 was transferred to Heteronychia Brauer &
Bergenstamm (Ezpierretia Rohdendorf) in an ad-
dendum of the original paper.)
The hypothesis that Sarcophaga amputata and S.
kunonis are sister species is corroborated by the
presence in these two species of at least five char-
acter states that are judged to be derived with regard
to the ground plan of Discachaeta and not found in
any other species of this taxon: 1) Postgena with at
least some black setae in anterior part, 2) Proanepi-
sternum setose, 3) Abdominal tergite 5 almost de-
void of microtomentum, 4) Male gonopod with
long setae on the median surface (fig. 3), and 5)
Female mid femoral organ reduced, i.e. indistinct or
absent. Sarcophaga amputata is easily separated
from S. kwnonis in both sexes by the greater exten-
sion of white postgenal setae. These cover at least
one half (strict lateral view) of the postgena in S.
amputata while they are almost absent in S. kuno-
nis. However, as the Madeiran fauna of Sarcopha-
gidae may still be incompletely known, the shape of
the aedeagus still provides the best clue to reliable
identifications.
Sarcophaga amputata may be widespread on the
main island and is the first species of Sarcophagidae
recorded from the Desertas islands.
Sarcophaga (Pierretia) iulicida sp.n. (figs. 5-9)
Type material. — Holotype 8, Portugal: Mouräo, ex
Ommatoiulus moreleti, larva 8.v.1987, pupa 14.v.1987,
adult 30.v.1987, P.T. Bailey (BMNH). The holotype is in
good condition, with the puparium glued to a piece of
cardboard and the dissected terminalia stored in glycerine
in a microvial, both items pinned with the specimen.
Description
Male. — Head. Narrowest part of frons 0.25 X
head width. Parafacial plate with a row of setae
along the eyemargin. The upper part of the row
consists of uniserial setae, the lower part is irreg-
ularly biserial with 3-5 long bristles anterior to the
x
PAPE: New species of Sarcophaga
setae. Arista with longest hairs about 4 x as long as
second aristomere. Gena with black setae only,
postgena with white setae only.
Thorax. Proanepisternum bare. Chaetotaxy: 2-3
distinct pairs of presutural acrostichals, postsutural
acrostichals (including prescutellars) not differen-
tiated, dorsocentrals = 3 + 3, intra-alars = 1 + 2,
supra-alars = 1 + 3, postalars = 2. Scutellum with
2 pairs of lateral scutellar bristles, 1 pair of short
apicals, and 1 pair of discals. Mid femur with 3
anteroventral bristles and a row of posteroventrals.
A pical posteroventrals slightly stronger than apical
anteroventrals. Hind tibia without elongated setae.
Abdomen. Tergites 3-4 with strong median mar-
ginals on T5 with a complete row of marginals.
Terminalia black, protandrial segment without
marginal bristles. Aedeagus with well developed
harpes, each of which has a narrow winglike pro-
cess that is directed laterally and opposite to its
counterpart. Vesica reduced to a simple swelling or
hump. Juxta moderately long and deeply cleft.
Length: 7 mm (estimated from dissected holo-
type).
Female. — Unknown.
Puparium of the usual barrel-shape and with the
posterior spiracles in a deep pit as in most species
of Sarcophaginae.
Distribution. — Palaearctic: Portugal.
Figs. 5-9. Sarcophaga tulicida, male ter-
minalia. 5, right cercus and surstylus,
lateral view, setae omitted; 6, cerci, pos-
terior view, setae omitted; 7, aedeagus,
lateral view; 8, distiphallus, dorsal (=
posterior) view; 9, right paramere +
gonopod, lateral view. Abbreviations: h
= harpes, j = juxta. Scale = 0.1 mm.
41
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Etymology. — A noun in apposition. From the
Latin wluys = multiped, millipede, and -cida = suffix
denoting cutter, killer, killing. The name refers to
the millipede-parasitizing habit of the species.
Remarks. — The present species belongs to a
probably monophyletic group whose members all
possess an aedeagus very similar to that of Sarco-
phaga nigriventris Meigen. The group is defined by
the characteristic and probably apomorphic shape
of the aedeagal juxta, which can be dissolved into
the two character states: 1) Juxta deeply cleft, and
2) Juxtal prongs tapering. Both states are readily
seen on figs. 7-8. No formal genus-group name has
been applied explicitly to this taxon, but the concept
of Pierretia Robineau-Desvoidy (sensu stricto) of
Verves (1986) is very close, the only difference
being that this author includes Sarcophaga granu-
lata Kramer, which does not possess either of the
derived character states. I prefer to restrict the
name Pierretia, as a subgenus of Sarcophaga, to the
group of species possessing the two above menti-
oned character states, ie. Sarcophaga discif-
era/Pandellé, S. sulicida, S. lunigera Böttcher, S.
nigriventris, S. socrus Rondani, $. soror Rondani, S.
sororcula (Rohdendorf) and S. villeneuvei Böttcher.
Sarcophaga tulicida may easily be separated from
other members of the subgenus by the wing-like
projection of each of the harpes which is directed
laterally and set at right angles to the longitudinal
axis of the aedeagus. This is most easily seen in
dorsal (= posterior) view (fig. 8).
The biology of Sarcophaga iulicida is interesting
as very few Diptera are known to parasitise milli-
pedes, and among the Sarcophagidae only species of
the New World genus Spirobolomyia Townsend
have been repeatedly bred from live millipedes. The
habits of other species of the subgenus Pierretia (as
defined above) indicate a broad to very broad spec-
trum of hosts or prey. Thus, the species Sarcophaga
nigriventris has been bred from snails, beetles,
grasshoppers and bees (see references in Pape
1987).
ACKNOWLEDGEMENTS
I extend my sincere thanks to Mr J.C. Deeming,
Cardiff, and Mr A.C. Pont and Mr N.P. Wyatt, both
London, for loan of specimens in their care.
REFERENCES
Downes, W. L., Jr., 1965. Family Sarcophagidae. Pp.
933-961. In: A. Stone et alii (eds): A catalog of the
Diptera of America north of Mexico. — Agriculture
Handbook No. 276, Washington, D.C.
Pape, T., 1986. The Sarcophagidae (Diptera) of Madeira,
with the description of a new species of Discachaeta
Enderlein. - Bocagiana 93: 1-4.
Pape, T., 1987. The Sarcophagidae (Diptera) of Fenno-
scandia and Denmark. — Fauna entomologia scandi-
navica 19: 1-203, 2 pls.
Pape, T., 1988. A revision of the Palaearctic Sarcophagidae
(Diptera) described by C. Rondani. - Stuttgarter Bei-
träge zur Naturkunde (Serie A) 416: 1-22.
Povolny, D., 1986. Discachaeta gigas sp.n. und Heterony-
chia (Heteronychia) vachai sp.n. - zwei unbekannte
Sarcophaginae-Arten aus dem Dinarischen Küstenge-
biet (Diptera, Sarcophagidae). — Acta Universitatis
Agriculturae 34(2): 229-238.
Séguy, E. 1941. Etudes sur les mouches parasites. 2. Cal-
liphoridae, calliphorines (suite), sarcophagines et rhi-
nophorines de l'Europe occidentale et méridionale. —
Encyclopédie entomologique (A) 21: 1-436.
Verves, Yu. G. 1986. Family Sarcophagidae. Pp. 58-193.
In: A. Soös (ed.): Catalogue of Palaearctic Diptera. Vol.
12, Calliphoridae - Sarcophagidae. - Akadémiai Kiadó,
Budapest.
Received: 29 May 1989
Accepted: 17 July 1989
THOMAS PAPE
Zoologisk Museum, Copenhagen
REVISIONARY NOTES ON AMERICAN
SARCOPHAGINAE (DIPTERA: SARCOPHAGIDAE)
Pape, T., 1990. Revisionary notes on American Sarcophaginae (Diptera: Sarcophagidae).
Tijdschrift voor Entomologie 133: 43-74, 66 figs. ISSN 0040-7496. Published 31 July 1990.
Selected New World genera of Sarcophaginae are redefined: Argoravinia Townsend, 1917
is treated as a senior synonym of Raviniopsis Townsend, 1918, syn. n., Comasarcophaga
Hall, 1931 as a senior synonym of Archimimus Reinhard, 1952, syn. n., and Emblemasoma
Aldrich, 1916 as a senior synonym of Pessoamyia Lopes, 1938, syn. n. and Colcondamyia
Reinhard, 1963, syn. n. Microcerella Macquart, 1851 is considered as synonymous to the tribe
Microcerellini, except that Cryptosarcophila Townsend, 1931 is excluded, and 6 new names
are proposed to replace junior secondary homonyms: Microcerella aulacophyto nom. nov.
for Aulacophyto auromaculata Townsend, 1919; Microcerella austrohartigia nom. nov. for
Austrohartigia bicoloricauda Lopes, 1981; Microcerella boettcherimima nom. nov. for
Boettcherimima hypopygialis Lopes, 1950; Microcerella boetia nom. nov. for Boettia ecua-
toriana Lopes, 1982; Microcerella carchia nom. nov. for Carchia andina Lopes, 1982; and
Microcerella jujuyia nom. nov. for Jujuyia alvarengai Lopes, 1980. Microcerella bermuda sp.
n. (Bermuda Is) and M. adelphe sp. n. (eastern Canada & USA) are described and the
phylogenetic relationships to other members of the genus are discussed. Microcerella
acridiorum (Weyenbergh, 1875) is considered a senior synonym of Xenoppiella dyscineti
Blanchard, 1966, syn. n. Fletcherimyia Townsend, 1917 and Spirobolomyia Townsend, 1917
are discussed in a phylogenetic context and it is argued that both taxa should be excluded
from Blaesoxipha (s.l.) |Protodexiini + Impariini]. Species limits within the genera are
revised and Fletcherimyia abdita sp. n. (southeastern USA) and Spirobolomyia latissima sp.
n. (southeastern USA & Mexico) are described. Tripanurga Brauer & Bergenstamm, 1891
is redefined and treated as a senior synonym of Metoposarcophaga Townsend, 1917, syn.
n. and Erwcophaga Reinhard, 1963, syn. n. Tripanurga guatemala sp. n. (Guatemala) is
described, and a probable sister group relationship to T. villipes (Wulp, 1895) is demon-
strated. Infraspecific variation within T. aurea (Townsend, 1917) is discussed and the species
is divided into a Baja Californian form and a mainland form.
Dr T. Pape, present address: Danish Bilharziasis Laboratory, Jaegersborg Allé 1 D, DK-2920
Charlottenlund.
Key words. — Diptera; Sarcophagidae; generic redefinitions; new species; New World.
CONTENTS INTRODUCTION
Imrtoductionsv een NUE ee 43 The New World fauna of Sarcophagidae deviates
Methods, terminology, depositories............ 44 markedly from that of the Old World with regard
Generic redefinitions and descriptions of to diversity. The subfamily Miltogrammatinae has
SPÉCIES Rae ne RL N or sdraiata 45 significantly fewer species, which in part is to be
GenustAncoraviniane RAMA 45 expected as the fauna of the Palaearctic Region is
GENUSUCO4ANEOPPAZARA IO 45 much more fully known and the extension of gener-
GeUStE7 IENA OMAR OR 46 ally species-rich areas as dry savannas, shrublands
Genussilerchenn na 47 and semi-deserts is much larger, e.g. on the African
Genus Vierocenell a RENE ER EE 48 mainland and in Central Asia. Within the subfamily
GENUSS piobolon pn 53 Sarcophaginae, on the other hand, species richness
GENIUS ANZIO 55 per se is greater in the New World, and as the
cknowledoemenesn O 58 sarcophagine fauna of especially the neotropics,
References TNA leeren. 58 where the large majority of species occur, is still
43
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
very insufficiently known, this difference may be-
come even more pronounced with time. The New
World fauna of Sarcophaginae has diversified
morphologically and biologically to a degree unri-
valled by the Old World members of this subfam-
ily. Some New World species have green eyes,
several have a metallic abdomen, and variations
in setosity and configuration of the male termina-
lia are legio and often much more radical than
those of the Old World species. The New World
species Peckia gulo and Tricharaea occidua repres-
ent the largest and the smallest species of Sarco-
phaginae, respectively, and sarcophagine life hab-
its include parasites of millipedes and insects, as
well as predators of reptile eggs, hymenopteran
kleptoparasites and generalized scavengers. The
much greater morphological and biological diver-
sity of New World species of Sarcophaginae com-
pared to those of the Old World has caused pro-
nounced differences of opinion regarding generic
limits within this subfamily (especially so, because
many of the previous authors have been more in-
terested in differences than in similarities). Nu-
merous monotypic genera have been proposed,
with no other warranty than their ‘uniqueness’
and/or weirdness.
My research on New World Sarcophaginae, espe-
cially in connection with a major conspectus of the
genus Blaesoxipha covering species limits and in-
ter- and infrageneric phylogenetic relationships
(Pape in prep.), during a one-year fellowship at the
National Museum of Natural History (USNM),
Washington, D.C., has led me to propose some
phylogenetically more consistent generic defini-
tions. It is felt that these definitions, in addition to
being theoretically sound, support identification
and information retrieval purposes better, and it is
considered appropriate to make the information
available as a single paper, with descriptions of
several hitherto unrecognized species which turned
up during my study, rather than split it into several
generic revisions and isolated descriptions of spe-
cies. Thus, it should be stressed that the present
paper is far from a comprehensive treatment of all
New World genera, and the fact that most genus-
group taxa have been excluded does not mean that
these are accepted by me. Older genus-group taxa
such as Pachygraphia Brauer & Bergenstamm,
1891, Lepidodexia Brauer & Bergenstamm, 1891,
and Johnsonia Coquillett, 1895 are especially in
need of a critical revision and will probably have to
be redefined as much broader entities in order to
absorb the numerous ‘less aberrant’ and probably
paraphyletic genera that often are considered
‘closely related’ to these genera. Much remains to be
done before generic limits stabilize, and more tho-
rough character analyses are needed before the taxa
above species level can emerge as well corroborated
monophyletic groups.
44
METHODS, TERMINOLOGY, DEPOSITORIES
Specimens were dissected by conventional cut-
ting of terminalia and heating in potassium or so-
dium hydroxide for about five minutes with subse-
quent rinsing in distilled water and transfer
through alcohol to glycerine. Drawings of termina-
lia were made from glycerine preparations, either
directly by using an ocular grid or from semiper-
manent slide mounts in glycerine gel using a pro-
jecting compound microscope.
Names of genus-group and species-group taxa
mentioned in the text are given without the con-
ventional citing of the original author except where
this has any direct bearing on nomenclature, e.g. in
the lists of synonyms. According to the Interna-
tional Code on Zoological Nomenclature, authors
do not form part of the name proper (ICZN: 51,a),
and they often serve no other purpose than as a
reference to the original source of the name. Recent
revisions are usually a better authority on the iden-
tity of a species or genus than the original descrip-
tion. For the present paper, identities of taxa not
explicitly defined are easily retrieved from the ref-
erences cited in the associated discussion or from
the most recent regional catalogues, i.e. Downes
(1965) and Lopes (1969), but see also Shewell
(087)
Morphological terms follow McAlpine (1981)
except that I have adopted Sabrosky’s (1983) use of
'microtomentum’ for the dense cuticular micro-pu-
bescence that may give surfaces a pollinose or prui-
nose appearance.
As no generic cladogram of the Sarcophaginae
has been produced in the present or any other paper
I have made no attempt of evaluating transforma-
tion series polarities for the character states used in
the generic diagnoses, i.e. whether apomorphic or
plesiomorphic at the level in question (which would
be nothing but ‘qualified guesses’ based on my own
more or less explicitly developed ideas of generic
phylogeny). The character states listed in the diag-
noses are, of course, only a fraction of those known,
and I have restricted the lists to contain only those
states that seem to be unique (and therefore possi-
bly autapomorphic) as well as states judged by me
to possess ‘diagnostic power’ (i.e., they will in com-
bination serve to define the taxon in question).
The institutions from which material examined
or otherwise referred to is deposited have been
abbreviated with the following acronyms:
AMNH: American Museum of Natural History,
New York, USA;
CAS: California Academy of Sciences, San
Francisco, USA;
CNC: Canadian National Collection of Ar-
thropods, Ottawa, Canada;
FSCA: Florida State Collection of Arthropods,
Gainesville, USA;
MACN: Museo Argentino de Ciencias Naturales,
Buenos Aires, Argentina;
MCZ: Museum of Comparative Zoology, Cam-
bridge, USA;
MNHN: Museum National d'Histoire Naturelle,
Paris, France;
NMW: Naturhistorisches Museum Wien, Austria.
UNAM: Instituto de Biologia, Universidad Na-
cional Autónomia de Mexico;
USNM: National Museum of Natural History,
Washington, D.C., USA;
ZML: Zoologiska Museet, Zoologiska Institu-
tionen, Lund, Sweden;
ZMUC: Zoological Museum, University of Co-
penhagen, Denmark.
GENERIC REDEFINITIONS AND DESCRIPTIONS OF
SPECIES
Genus Argoravinia
Argoravinia Townsend, 1917b: 190, 193, 195. Type spe-
cies: Sarcophaga argentea Townsend, 1912 (= Sarco-
phaga rufiventris Wiedemann, 1830), by original de-
signation.
Raviniopsis Townsend, 1918: 160. Type species: Ravint-
opsis aurea Townsend, 1918, by original designation.
Syn. n.
Sarcophagina Curran, 1928: 102. Type species: Sarcopha-
gina candida Curran, 1928, by original designation.
Pachygraphomyia Hall, 1933: 259. Type species: Pachy-
graphomyia spinosa Hall, 1933, by original designa-
tion.
Diagnosis. — The genus Argoravinia may be
defined by the following character states:
1) Aedeagus with a very long and S-shaped me-
dian stylus, see, e.g., Roback (1954: fig. 39);
2) One or more setae on the underside of the stem
of r,,, elongate;
3) Male cercal prong bent backwards;
3) .Ejaculatory apodeme large.
As tentatively suggested by Pape (in press), the
genera Argoravinia and Raviniopsis should be
treated as synonyms. They have been included in a
subtribe of their own (within Sarcodexiini) by
Lopes (1975a, 1982e), the monophyly of which al-
though never corroborated by explicit reference to
probable apomorphies has never been questioned.
Lopes (1982e) lists a number of character states, but
only some of these can be considered derived with
regard to the groundplan of the Sarcophaginae, and
this author does not propose any most probable
sister group. The two very distinctive character
states 1-2 mentioned above seem to be autapomor-
phic and will fully define the genus. The remaining
two character states may diagnose the taxon
further, but these may be plesiomorphic at the level
of Argoravinia (i.e. define a more inclusive group)
as discussed under Tripanurga.
PAPE: American Sarcophaginae
Having corroborated the monophyly of Argora-
vinia, it remains to be discussed why a splitting is
opposed. One argument is the small size of this
taxon, with a total of 7 species recognized at pres-
ent. Even if monophyletic subgroups can be recog-
nized (and in theory a complete resolution into
dichotomies should be possible), nothing is gained
by erecting formal genus-group names if one then
has to create new family-group names (e.g. tribes
and subtribes) to keep the related taxa together.
Another, more formal, argument is the subgroups
proper. Argoravinia in the narrow sense of Lopes
(1976), including A. rufiventris and A. alvarengat,
is most probably monophyletic, defined by the aut-
apomorphic process from the posteroventral mar-
gin of the epandrium that partially covers the sur-
stylus, and by the much more distinctly S-shaped
median stylus compared to the remaining Argora-
vinta (see figs. in Lopes 1976 and 1988b). I have not
found any character state shared only by these re-
maining species, which were treated as Raviniops
by Lopes (1988b), that may be considered apomor-
phic at this level, and none have been suggested. If
the monophyly of the taxon cannot be corroborated,
it has to be either split or widened until fitting a
pattern for which character state distributions cor-
roborate a hypothesis of strict monophyly. In the
present case, the obvious choice is a merging with
the older Argoravinia.
Genus Comasarcophaga
Comasarcophaga Hall, 1931: 280. Type species: Comasar-
cophaga texana Hall, 1931, by original designation.
Tejasomyia Reinhard, 1945: 68. Type species: Tejasomyta
nexilis Reinhard, 1945, by original designation.
Archimimus Reinhard, 1952: 140. Type species: Archimz-
mus camatus Reinhard, 1952, by original designation.
Syn. n.
Diagnosis. — The genus Comasarcophaga may
be defined by the following character states:
1) Male mid femur apically with a short comb-like
row of posteroventral spines;
) Male cercal prong bent backwards;
) Aedeagal juxta slightly displaced ventrally rel-
ative to longitudinal axis of aedeagal tube, the
latter thereby appearing 'hump-backed' (figs.
1-4, additional figs. in Lopes 1979 and Lopes &
Tibana 1988);
4) Aedeagal vesica appearing more or less square
in lateral view (cf. figs. mentioned above).
D N
The genus Comasarcophaga was described by
Hall (1931) as a monotypic genus for his new spe-
cies C. texana. Downes (1965) synonymized Coma-
sarcophaga with Tejasomyia, thereby adding the
species C. nexilis, and apparently only these two
species were included in the genus by Shewell
(1987: 1168), who gave "2 spp.; Texas, California”.
Another species, Sarcophaga prolepsis, described
45
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
by Reinhard (1947) from Arizona, was transferred
by Downes (1965) to Blaesoxipha subgenus Servai-
sia, and later by Lopes (1988d) to Archimimus.
Lopes & Tibana (1988: 324) defined Archimimus in
part by the features: "styli of glans and median
process spinous” (lateral and median styli in my
terminology) and "arista plumose to tip”. However,
by including prolepsis and longespinus (Lopes
1988d), which both possess simple lateral and me-
dian styli with small denticles as well as a pubescent
or short-plumose arista with a distinctly bare tip,
conditions perfectly like those found in most other
Sarcophaginae, the definition breaks down at least
for these characters. Other character states listed by
Lopes & Tibana (1988) as diagnostic for Archimz-
mus are: 3-Á post-dorsocentral bristles, proanepis-
ternum bare, male mid femur with apical comb-like
posteroventral spines, aedeagal vesica strongly
sclerotized, and female tergite 8 (7 in my terminol-
ogy) setose. These states, however, do not secure an
unambiguous definition of the taxon as this combi-
nation is found in other genera, e.g. Blaesoxipha in
part and Comasarcophaga.
Of the four states listed in the definition above,
1 and 2 are found in several other genera. Only 3
and 4 seem to be unique (i.e., autapomorphic), but,
unfortunately, they are less clear cut and more dif-
ficult to describe as are many other characters of the
male terminalia. Comasarcophaga in the present
definition may be divided into two groups based on
similarity (as perceived by me) of the male termi-
nalia: one containing longispinus, nexilis, prolepsis
and texana, another containing the remaining
members. This agrees with Lopes (1988d: 918),
who mentioned that longispinus and prolepsis was
“provisionally included in Archimimus in spite of
some important differences from the type-species:
last female sternite large and clypeal arch of the
first instar larva complete’. Differences as such,
however, cannot reject phylogenetic relationships,
and as I have found no better way to describe the
species explicitly in terms of shared character
states, lumping all Archimimus with nexilis and
texana in a broader Comasarcophaga is necessary to
maintain the monophyly of the taxon.
Terminalia of species of Comasarcophaga not
figured by Lopes (1979) and Lopes & Tibana (1988)
have been illustrated in figs. 1-12.
The only biological information available for Co-
masarcophaga is a single breeding record of a male
C. texana from the bulimulid snail Rabdotus deal-
batus, and the fact that another male of this species
has been collected from a dysentery fly trap (Neck
& Lopes 1973).
Genus Emblemasoma
Emblemasoma Aldrich, 1916: 56. Type species: Emble-
masoma erro Aldrich, 1916, by original designation.
46
Pessoamyia Lopes, 1938: 333. Type species: Pessoamyia
prosternalis Lopes, 1938, by original designation. Syn.
n.
Colcondamyia Reinhard, 1963a: 82. Type species: Colcon-
damyia falcifera Reinhard, 1963a, by original designa-
tion. Syn. n.
Golcondamyia Reinhard, 1963b: 152. Unjustified emenda-
tion of Colcondamyia.
Diagnosis. — The genus Emblemasoma may be
defined by the following character states:
1) Prosternum broadened (Shewell 1987: fig. 28);
2) Male mid femur apically with a row of comb-
like posteroventral spines.
When Lopes (1938) described Pessoamyia he did
not present any hypothesis as to possible relation-
ships to other genera, and the main reason for
describing the genus apparently was the "proster-
num extraordinariamente desenvolvido” (p. 333).
Later, Lopes (197 1b: 89) presented a "study of spe-
cies showing large and inflated prosternum”, al-
though without commenting on the similarly
equipped Colcondamyia, and he uncritically ac-
cepted the validity of the two genera discussed (ie.
Emblemasoma and Pessoamyta). Actually, he did
not even mention whether or not he considered the
enlarged prosternum as indicating phylogenetic re-
lationship, but he later erected the subtribe Emble-
masomatina (Lopes 1975b, as tribe in 1982e and
1988c), including all the above-mentioned genera
(Lopes 1982e), and he explicitly recognized the
modified prosternum to be autapomorphic within
the Sarcophagidae (Lopes 1988c).
As shown in the list of synonyms given above, I
prefer to reduce the (sub)tribe to the generic level,
and at least three reasons should be considered.
Firstly, the comparatively small size of the entire
taxon (i.e. Emblemasoma in the present sense),
does not, in my opinion, necessitate a generic sub-
division to enhance the clarity of the phylogenetic
relationships between the included species. Admit-
tedly, a group consisting of Pessoamyia and Em-
blemasoma sensu Lopes (1988c) may probably be
monophyletic based on the shared possession of a
highly modified male cercus, which in profile has a
subapical dorsal knob and ends in a blunt hook
(Lopes 1988c: figs. 17, 42, 45), but I prefer to name
this assemblage of species the erro species-group
rather than use a formal genus-group name. Se-
condly, while the monophyly of Emblemasoma in
the present sense seems reasonably well corrobo-
rated (the species are also biologically similar in
being parasites or predators of insects, as far as
known), the monophyly of the groups recognized
by Lopes (1988c) seems much less corroborated.
Thus, the only derived character states defining the
taxon Pessoamyia seem to pertain to the aedeagus
(vesica reduced, vesica spinous, ‘lateral plates’ hya-
line, see key in Lopes 1975b), and Lopes (1988c: 17)
admitted that males are needed to decide whether
species for which only females are known “belong
to Pessoamyia, Emblemasoma or to a distinct ge-
nus’. Thirdly, the enlarged prosternum provides an
easy-to-see feature for the recognition of the taxon
and thus of value from the point of view of the non-
specialist (e.g. when using a generic key).
Note that Pape (1987) included Emblemasoma
auditrix in a wide concept of Blaesoxipha, and the
bent male cercus and the fused abdominal sternites
6-8 of auditrix are indeed reminiscent of the condi-
tion found in some species of Blaesoxipha (Soper et
al. 1976). However, my revised concept of the latter
genus (Pape in prep.) includes all sarcophagine
species with the lateral styli non-conducting and
fused ventrally through a median sclerotization
(?vesica). Therefore, the three free and apparently
sperm-conducting (i.e. hollow and communicating
with the spermduct) styli of Emblemasoma com-
bined with the lack of a row of trochanteral spines
provides evidence that neither E. audıtrıx nor all of
Emblemasoma should be included in Blaesoxipha
(see also discussion under F/etcherimyia and Spiro-
bolomyia). Evidence for either placement of audz-
trix, however, is rather sparse, and I prefer the
present inclusion of this species in Emblemasoma
mainly because an inclusion within Blaesoxipha
would require that the free styli should be consi-
dered a reversal, which so far has not been shown
to occur in any other species of Sarcophaginae. A
detailed numeric cladistic analysis of all sarcopha-
gine genera is needed to test this hypothesis.
Genus Fletcherimyia
Fletcherimyia Townsend, 1917b: 191. Type species: Sar-
cophaga fletcheri Aldrich, 1916, by original designa-
tion.
Peltopyga Townsend, 1917b: 191, 194. Type species: Sar-
cophaga celarata Aldrich, 1916, by original designation.
Diagnosis. — The genus Fletcherimyia may be
defined by the following character states:
1) Male mid femur apically with a short comb-like
row of posteroventral spines;
2) Male cercal prong bent backwards;
Aedeagus with lateral styli shortened;
4) Aedeagal vesica a single tongue-shaped struc-
ture (figs. 13-18);
5) Aedeagal juxta with cuticular pubescence along
the distal margin (figs. 13-18);
6) Female abdominal tergite 6 strongly convex;
7) Female abdominal sternites 6-7 fused;
Larvae unable to develop to maturity outside
pitchers of Sarracenia.
Character states 5 and 6 seem to be autapomor-
phic.
Fletcherimyia and Peltopyga, both monotypic,
were described in the same paper by Townsend
PAPE: American Sarcophaginae
(1917b). When Townsend (1938) later redescribed
these genera, he still included only the type species
in Peltopyga while he did not state how many spe-
cies were included in F/etcherimyia. Downes (1965)
catalogued the Nearctic species and treated Fletche-
rimyia and Peltopyga as synonyms, including four
similar species, all of which were known to deposit
their larvae in pitchers of Sarracenia, in Fletcheri-
myia as subgenus of Blaesoxipha.
This apparently obligate association with pitcher
plants is reflected in the distribution in that all
species of Fletcherimyia have their center of distri-
bution in the eastern part of the continent. Buckell
& Spencer (1957) listed Sarcophaga fletcheri from
Canada (Robson, B.C.), but I have been unable to
confirm this and the record was apparently not
accepted by Downes (1965). Species of Sarracenia
are mainly eastern in distribution although the
most wide-ranging species, $. purpurea, reaches far
west to the District of Mackenzie and the extreme
northeastern part of British Columbia (Cody & Tal-
bot 1973). Sarracenia purpurea is known from one
locality only in B.C., a peat bog approximately 30
km south of Fort Nelson (Krajina 1968), and the
record of fletcheri from Robson seems unlikely. It
should be expected that species of Fletcherimyia
require a certain density of Sarracenia localities to
be able to maintain stable populations.
Males of Fletcherimyta can often be seen resting
on the rim of the Sarracenia pitchers, and these may
serve as the aggregation site. Females larviposit
large first stage larvae singly into newly opened
pitchers, and the larvae feed on invertebrates
caught in the fluid. The larva lives submerged, but
with the posterior end kept at the water surface to
facilitate respiration. The mature larva leaves the
pitcher and pupates at the base of the plant (For-
syth & Robertson 1975, Fish 1976, Fish & Hall
1978).
A few other species of Sarcophagidae breed in the
pitchers of insectivorous plants: The Nearctic Sar-
cophaga (Liosarcophaga) sarraceniae in Sarracenia
pitchers (Aldrich 1916) and the Oriental Sarco-
phaga (Pierretia) urceola in Nepenthes pitchers
(Beaver 1979, Shinonaga & Beaver 1979), but these
are easily shown by morphological evidence to have
evolved the life habit independently from species of
Fletcherimyia, and even from each other, as indi-
cated by the subgeneric assignment given in paren-
theses.
Roback (1954) gave figures of the aedeagus of the
four species known to him, but instead of straight-
ening out the homologies of the acrophallic sclero-
tizations, he simply introduced the morphological
term 'stematis’ for the type of acrophallus found in
Fletcherimyia. Lopes (1971a: 4) stated that "the
opening [of the acrophallus] is very much compli-
cated” and proposed that it had “a different origin”
from that of Blaesoxipha. This, however, seems
47
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
little warranted as there is no evidence that the
sperm exit proper is not homologous throughout
the Sarcophagidae (and probably all of Diptera),
and therefore may serve as an important morpho-
logical landmark. If this is accepted, Fletcherimyia
falls nicely within the groundplan pattern of the
Sarcophaginae in that all species possess three styli
which all seem to be more or less tubular and con-
nected to the common sperm duct (fig. 18).
With the additional species described below, the
genus contains five species, but some other species
should be mentioned as they have been assigned to
the genus Fletcherimyia. Lopes (1946) described F.
speciosa from Mexico and included Sarcophaga ces-
sator as well, and Rohdendorf (1971) described
Fletcherimyia zayasi from Cuba. These three spe-
cies, however, all belong to Blaesoxipha, the mo-
nophyly of which is corroborated mainly by the
transformation of the lateral styli into non-conduct-
ing structures and a sclerotized ventromedian
bridge connecting these (Pape unpubl.). It should
be stressed that although Lopes (1971a: 5) stated
that Fletcherimyia (and some other taxa) did "not
belong to Tephromyiini |= the present Blaesoxi-
pha in part] simply because they were "good
genera”, it is perfectly possible to argue for the
inclusion of Fletcherimyia within Blaesoxipha, e.g.
as a subgenus (Downes 1965), simply by widening
the concept of Blaesoxipha sufficiently (e.g. by de-
fining this genus on the comb-like pv bristles of the
male mid femur). This does not, of course, alter the
well corroborated hypothesis that species of Flet-
cherimyia as presently recognized, form a mono-
phyletic group. My reason for not including Flet-
cherimyia within Blaesoxipha is primarily that the
resulting taxon, although possibly monophyletic if
also containing Comasarcophaga and Spirobolo-
myia discussed elsewhere in the present paper,
would be much more vaguely defined in the sense
that the monophyly would be much more tentative.
Considering the well corroborated monophyly of
Blaesoxipha, Comasarcophaga, Fletcherimyia and
Spirobolomyia respectively, as well as their rather
different external morphology, I see no convincing
arguments for combining them into one taxon. It
could be argued that the combined taxon would be
more easy to identify in conventional sorting of
specimens, as dissections of the male terminalia,
which often are necessary for the non-specialist and
therefore prevent handling a large number of spe-
cimens, would be reduced to a minimum. I still
prefer, however, to give the criterion of monophyly
higher priority than that of usefulness.
Fletcherimyia abdita sp. n. (figs. 13, 19, 24)
Type material. — Holotype 4, USA: Alabama, Theo-
dore, reared from Sarracenia drummondii, 10.vi.1916,
48
FM. Jones [paratype of Sarcophaga rileyi, cf. Aldrich
(1916: 243)] (USNM). Paratypes. - USA: 14, data as
holotype but with the date 22.vi.1916 [paratype of Sarco-
phaga rileyi) (USNM); 26, locality as holotype but with-
out date (FSCA, ZMUC); Florida, Freeport, 26,
25.v1.1921, [no collector, label in Jones’ handwriting],
reared from Sarracenia drummondii (FSCA); Mississippi,
Biloxi, 18, 6.vi.1921, 28, 23.vi.1921, 28, 25.vi.1921, FM.
Jones, reared from Sarracenia sledgei (4 in FSCA, 1 in
ZMUC).
Description
Male. — General morphology similar to other
species of Fletcherimyia (see descriptions in Al-
drich 1916), but separated from these by the struc-
ture of male abdominal sternite 5 and male termi-
nalia (compare figs. 13-28). The following
description is restricted to these diagnostic features.
Abdomen: Posterior margin of sternite 5 with
accessory lobes like those of F. celarata but slightly
raised from the plane of the remaining sternite so
that a small flange penetrates behind (morpholo-
gically above) it (fig. 19). Sternite 5 of F. celarata has
accessory lobes broadly rounded, level with remain-
ing sternite (fig. 20), and dark brown along poster-
ior margin in contrast to the yellowish remaining
sternite. Sternite 5 of F. rileyt has tapering accessory
lobes and posterior margin concolorous with re-
maining sternite (fig. 23).
Terminalia: Cercus, as seen in posterior view,
with an outline intermediate between that of F.
fletcheri and F rileyi. The aedeagus is similar to that
of E fletcheri, but the juxta is broader and more
square and the vesica is larger with a spinelike
process at about middle (fig. 13).
Female. — Unknown.
Etymology. — A Latin adjective, from abditus =
hidden, concealed, put away. The name refers to the
fact that the holotype was found amongst paratypes
of F. rileyi.
Distribution. — Nearctic: USA (Alabama, Flor-
ida, Mississippi).
Biology. — Larvae scavengers in pitchers of Sar-
racenia drummondu and S. sledget.
Genus Microcerella
Microcerella Macquart, 1851: 209 (236). Type species:
Microcerella rufomaculata Macquart, 1851, by original
designation.
Doringia Weyenbergh, 1875: 85. Type species: Nemorea
acridiorum Weyenbergh, 1875, by original designation.
Syn. n.
Euparaphyto Townsend, 1912: 359. Type species: Expara-
phyto alpina Townsend, 1912, by original designation.
Syn. n.
Xenoppia Townsend, 1915: 20. Type species: Xenoppta
hypopygialis Townsend, 1915, by original designation.
Camptopyga Aldrich, 1916: 41. Type species: Camptopyga
aristata Aldrich, 1916 [= Xenoppia hypopygialis
Townsend, 1915], by original designation.
Hypopelta Aldrich, 1916: 49. Type species: Hypopelta
scrofa Aldrich, 1916, by original designation.
Aulacophyto Townsend, 1919: 158. Type species: Aulaco-
phyto auromaculata Townsend, 1919 [secondary junior
homonym of Euparaphyto auromaculata Townsend,
1919: 157] [= Microcerella aulacophyto nom. nov.|,
by original designation. Syn. n.
Gymnopsoa Townsend, 1919: 161. Type species: Gym-
nopsoa texana Townsend, 1919, by original designa-
tion.
Catheteronychia Townsend, 1927: 230. Type species: Cat-
heteronychia chaetosa Townsend, 1927, by original
designation. Syn. n.
Xanthobrachycoma Townsend, 1927: 232. Type species:
Xanthobrachycoma analis Townsend, 1927, by original
designation. Syn. n.
Townsendmyia Prado & Fonseca, 1932: 167. Type species:
Stephanostoma argenteum Prado & Fonseca, 1932 |=
Sarcophaga (Bellieria) halli Engel, 1931], by original
designation. Syn. n.
Alaccoprosopa Townsend, 1934: 202. Type species: Alac-
coprosopa apicalis Townsend, 1934, by original desig-
nation. Syn. n.
Austrohartigia Townsend, 1937a: 115. Type species: Aus-
trohartigia magellanica Townsend, 1937a [= Sarco-
phaga spinigena Rondani, 1846], by original designa-
tion. Syn. n.
Microcerellamyia Hall, 1937: 354. Type species: Microcer-
ellamyia retusa Hall, 1937, by original designation.
Syn. n.
Itiophyto Hall, 1937: 355. Type species: Itiophyto engeli
Hall, 1937, by original designation. Syn. n.
Mimophytomyia Hall, 1937: 357. Type species: Mzmo-
phytomyia chilensis Hall, 1937 |= Alaccoprosopa api-
calis Townsend, 1934], by original designation.
Phaesarcodexia Hall, 1937: 362. Type species: Phaesarco-
dexia aldrichi Hall, 1937 (also as Brachicoma aldricht)
[= Sarcophaga spinigena Rondani, 1864], by original
designation.
Eumicrocerella Hall, 1938: 253. Type species: Eumicrocer-
ella duca Hall, 1938, by original designation. Syn. n.
Scopaediscus Blanchard, 1939: 802. Type species: Scopae-
discus muebni Blanchard, 1939, by original designa-
tion. Syn. n.
Boettcherimima Lopes, 1950: 706. Type species:
Boettcherimima hypopygialis Lopes, 1950 [secondary
junior homonym of Xenoppia hypopygialis Town-
send, 1915] [= Microcerella boettcherimima nom.
nov.], by original designation. Syn. n.
Parabypopelta Blanchard, 1955: 30. Type species: Parahy-
popelta salavini Blanchard, 1955, by original designa-
tion. Syn. n.
Chapiniola Dodge, 1965: 253. Type species: Chapiniola
impressa Dodge, 1965, by original designation. Syn. n.
Steatopyga Dodge, 1965: 254. Type species: Steatopyga
mirabilis Dodge, 1965, by original designation. Syn. n.
Boetia Dodge, 1965: 255. Type species: Boetia curiosa
Dodge, 1965, by original designation. Syn. n.
Xenoppiella Blanchard, 1966: 182. Type species: Xeno-
piella dyscineti Blanchard, 1966 |= Nemorea acridio-
rum Weyenbergh, 1875, syn. n.], by monotypy. Syn. n.
Azuayia Dodge, 1967: 681. Type species: Azuayıa tripar-
tita Dodge, 1967, by original designation. Syn. n.
Borgmeterisca Lopes, 1972: 349. Type species: Borgmeier-
PAPE: American Sarcophaginae
isca pilicoxa Lopes, 1972, by original designation. Syn.
n.
Townsendisca Lopes, 1974: 193. Type species: Townsen-
disca matucanensis Lopes, 1974, by original designa-
tion. Syn. n.
Xenoppina Lopes, 1975d: 573. Type species: Xenoppina
andina Lopes 1975d, by original designation. Syn. n.
Aulacophytoides Lopes, 1978a: 765. Type species: Az/aco-
phytoides alvarengai Lopes, 1978a, by original desig-
nation. Syn. n.
Jujuyia Lopes, 1980: 238. Type species: Jujuyia alvarengat
Lopes, 1980 [secondary junior homonym of Az/aco-
phytoides alvarengai Lopes, 1978a] [= Microcerella
jujuyia nom. nov.], by original designation. Syn. n.
Mallochisca Lopes, 1982b: 364. Type species: Mimophy-
tomyia mallochi Hall, 1937, by original designation.
Syn. n.
Penaisca Lopes, 1982b: 365. Type species: Penaisca qui-
maliensis Lopes, 1982b; by original designation. Syn.
n.
Cuzcomyia Lopes, 1982b: 367. Type species: Cuzcomyta
rufipes Lopes, 1982b, by original designation. Syn. n.
Carchia Lopes, 1982b: 368. Type species: Carchia andina
Lopes, 1982b [secondary junior homonym of Xenop-
pina andina Lopes, 1975d; [ Microcerella carchia nom.
nov.], by original designation. Syn. n.
Azuaya: Lopes (1982b: 363); erroneous subsequent spell-
ing of Azuayıa.
Boettia: Lopes (1982b: 363, 1982d: 607); erroneous sub-
sequent spelling of Boetza.
Parabybopelta: Lopes (1969a: 8, 81); erroneous subse-
quent spelling of Parahypopelta.
Phaeosarcodexia: Lopes (1969a: 8, 82; 1981: 327); errone-
ous subsequent spelling of Phaesarcodexia.
Townsendimyia: Lopes (1969a: 9, 1974: 195, 1975a: 271,
1982b: 363); erroneous subsequent spelling of Town-
sendmyia.
Diagnosis. — The genus Microcerella may be
defined by the following character states:
1) Eyes green (live or fresh material);
2) Male protandrial segment black;
3) Hypandrial arm swollen at level of gonopod;
4) Postgena with at least some black setae close to
genal suture.
The generic synonymizations presented above
are straightforward in the sense that the identities
of the type species of most genus-group names are
well known and the monophyly of what is here
considered as Microcerella has never been questi-
oned. One exception is the taxon Xenoptella dys-
cineti, which was mentioned as being a potential
member of the group by Lopes (1982b) but left with
no further comment, as the holotype (which is the
only specimen known) was not examined. This
uncertainty resulted from the insufficient original
description by Blanchard (1966), who gave a not
very informative description of the male terminalia
and did not provide any figures, only giving some
chaetotaxic characters and stating that the termina-
lia were "muy abultado” (p. 183). I have examined
the male holotype (MACN), which was mounted
on cardboard and completely hidden in dense
49
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
mould. After partly cleaning of the specimen and
remounting of the terminalia, it turned out to be a
typical member of Microcerella (s.l.) and a junior
synonym of M. acridiorum, syn. n. | Note that I have
not revised the type material pertaining to the lat-
ter name, and I have followed Lopes (1969b), who
reviewed its taxonomic history.] To corroborate
this, and to facilitate recognition and identification,
figures of the terminalia of the holotype of X. dys-
cineti are provided (figs. 29-32, compare Lopes
1969b: figs. 55-58). The holotype, which is perfectly
intact except for dissected (but well preserved) ter-
minalia, is heavily moulded and glued to a piece of
cardboard on its right side. Terminalia and an
empty puparium are mounted on separate pieces of
cardboard pinned with the specimen. It is labeled
"S/ Dyscinetus gagates, adulto.”, "Tandil 6-XI-1958
Bazan, R..”, "Xenopiella dyscineti ‚gen. et sp. nov.
det. E. E. Blanchard”, and "Xenopiella dyscineti E.
E. Blanchard 1966 Holotypus [red label]”.
Lopes (1982b) noted that the monotypic genus
Sarcophagulopsis (type species: S. trigonophymt)
possibly belonged to the Microcerellini. It is, how-
ever, a Blaesoxipha sensu lato and will be revised by
Pape (in prep.).
Somewhat ironically, the identity of the type spe-
cies of the genus Microcerella proper, M. rufomac-
ulata, has never been thoroughly revised. Séguy
(1925: 184) and Townsend (1931: 72, 1937b: 202)
examined and redescribed the female holotype in
MNHN, leaving no doubt of the assignment to
Microcerella in the present sense, but no attempt
has ever been made to associate the female with
male specimens, for which reason the identity is
highly uncertain. As other species are known from
the male sex only, it is very probable that M. rufo-
maculata is a senior synonym of one of these [an
obvious candidate being M. sarcophagina, the male
holotype of which, however, may be lost, as men-
tioned below |.
To accommodate all species of the taxa synony-
mized within Mzcrocerella above, a number of new
names have been proposed. These names, which all
should be treated as nouns in apposition, are
formed by using the name of their former genus as
species epithet. Two other secondary homonyms
are not covered in the list of generic synonyms.
They are given below, following the procedure
given above:
Microcerella boetia nom. nov. for Boettia [sic!]
ecuatoriana Lopes, 1982d: 613 [secondary junior
homonym of Euparaphyto ecuatoriana Lopes,
1982c: 495].
Microcerella austrohartigia nom. nov. for Austro-
hartigia bicoloricauda Lopes, 1981: 334 [secondary
junior homonym of Mesothyrsia bicoloricauda En-
derlein 019287151]:
50
The genus Mrcrocerella is defined in the most
narrow sense by Lopes (1969a) in the Neotropical
catalogue, as he includes only one species, M. sar-
cophagina, besides the type species M. rufomacu-
lata. The taxon M. sarcophagina, like M. rufomac-
ulata, has remained enigmatic since its description
by Thomson (1869), and the male holotype, which
is the only specimen known, seems to be lost. [P. I.
Persson (NMRS) has informed me that he has been
unable to recover the holotype, which was sent as
a loan to the late H. R. Dodge in the 1970’s, and I
did not succeed in finding it among the remnants
of the Dodge-collection in FSCA. | Without any clue
as to the structure of the terminalia, especially the
aedeagus, the identity can only be a qualified guess
whenever a detailed generic revision with extensive
information of species present in the type locality
concerned becomes available.
Lopes (1982b) provides a key to the genera of
Microcerellini, and his Microcerella keys out mainly
on the length of the first flagellomere relative to
that of the pedicel, i.e. whether the former is “twice
the length” (Microcerella) or only “a little longer”
(Microcerellamyia) than the latter, and his meas-
urements are apparently based partly on the ori-
ginal figures of Macquart (1851, pl. 22: fig. 1). It
seems to me, therefore, that although Microcerella
rufomaculata fits well within the tribe Microcerel-
lini, and thus within Mzcrocerella in the present
sense, the narrow concept of Microcerella sensu
Lopes needs revision and is tentative at best.
Lopes (1982b) discussed the limits of Microcer-
ellini and mentioned Austrobartigia, Doringia, and
Aulacophyto with ten, four, and five species respec-
tively (che latter now with an additional three des-
cribed by Tibana & Lopes (1988)), and he noted that
“the majority of the remaining [27] genera are
monotypic’ (p. 359). Note that Lopes (1982b) di-
vided the Microcerellini into the subtribes Micro-
cerellina and Hypopeltina. A single two-state char-
acter separates these two groups, namely the
possession of a pubescent versus a plumose arista
(key entry only), but obviously both states cannot be
assumed a priori to be apomorphic at this level as
no third state from which they could be derived
independently exists within the Sarcophaginae. No
further explanation is given and the classification
appears to be artificial. I have not been able to split
the tribe into a few well defined and probably mo-
nophyletic groups, and instead of accepting the
wealth of monotypic or very small genera that do
not convey much information (and actually often
are a burden for identification and information re-
trieval), I have lumped all species into one genus.
Downes (1965) and Shewell (1987), who are deal-
ing exclusively with the much sparser Nearctic
fauna, also apply a broader concept when they in-
clude four species within Mzcrocerella, one of which
has been recorded from the neotropical region (M.
hypopygialis, see Lopes 1980, 1982a). Microcerella
in this broadened sense is very well defined and
equals the tribe Microcerellini as presented in
Lopes (1982b, 1982e), except that I exclude the
monotypic genus Cryptosarcophila as discussed
below. Lopes (1969a: 7, 1982b: 361) mentioned that
all species seen by him in fresh condition have
green eyes, truly a unique condition within the Sar-
cophagidae. The green colour disappears after
death although the eyes may retain a feeble green-
ish tinge (at least for some years). I have not seen
live or fresh specimens of any of the Nearctic spe-
cies, and although at least M. texana seems to be
green-eyed (as judged from recently captured spe-
cimens in the USNM), information is needed from
living or freshly killed material. Therefore, al-
though I know of no exceptions within Mzcrocer-
ella, or of any (homoplasic) occurrences of green
eyes in other sarcophagine genera, I cannot fully
assess the strength of this character state for defin-
ing the genus. The majority of species of Mzcrocer-
ella have a bare, pubescent or short plumose arista
and males of almost all species have a dark brown
or black protandrial segment (first genital segment
of authors, syntergosternite 7-8 of McAlpine 1981),
distinctly contrasting to a bright red or orangish
epandrium. No species have their male terminalia
entirely red, and the black/black condition found in
M. muebni and in the probably monophyletic clade
LM. hypopygialis + M. texana + M. valgata] is most
probably derived from the black/red condition. The
aedeagus is highly characteristic by the very rigid
connection between basi- and distiphallus that
probably allows little or no articulation. A part from
a few species that have a highly modified aedeagus,
the ventral surface of the distiphallus is equipped
with a flat, sclerotized scale or plate. All species
seen by me have the hypandrial arms more or less
swollen at the level of the gonopods. The female
terminalia show fused sternites 6-8 much like the
condition seen in Spirobolomyia, Comasarcophaga,
and many Blaesoxipha, and tergite 6 is entire and
more or less extended beyond the posterior rim of
tergite 5.
Cryptosarcophila, with C. chaetosa as type spe-
cies and only species included, was transferred to
the Microcerellini by Lopes (1982b), but I prefer
not to include this taxon in Microcerella. The rea-
son for this is simply that I have not been able to
recognize any convincing shared and assumed syn-
apomorphic character state defining a clade con-
taining only Cryptosarcophila and Microcerella. Al-
though I have not examined any specimens of C.
chaetosa myself, Lopes (1975e: 46) redescribed the
species from the male holotype and noted that it has
the "genital segments red”, which most probably is
plesiomorphic relative to the red/black or black/
black condition in all Microcerella. Moreover, the
arista is plumose and the aedeagus has no scalelike
PAPE: American Sarcophaginae
ventromedian plate or process. Cryptosarcophila
chaetosa resembles many Microcerella in having
the white postgenal setae restricted to the hypos-
tomal bridge below the neck, which probably is a
derived condition relative to the groundplan of the
Sarcophaginae. Many other genera, however, seem
to share this character state, e.g. Johnsonia, Neo-
phyto and perhaps Notochaeta, and it does not
provide evidence for inclusion particularly in Mz-
crocerella.
Little information is available on the biology of
Microcerella, but species have been bred from land
snails, beetles and grasshoppers, and in the labor-
atory on horse blood serum (Blanchard 1966; Lopes
1969a, 1973).
Microcerella is most diverse in the neotropical
region and especially at high altitudes and in the
southern temperate zones. It was therefore surpris-
ing to find two specimens of an undescribed species
of Microcerella from Bermuda, and even more so as
the fauna of Sarcophagidae of these islands other-
wise seems to consist entirely of a few widespread
and possibly introduced species. Actually, the Dip-
tera fauna of the Bermuda Islands is rather meager
and with only few endemics. Many older Bermudan
records of Sarcophagidae are misidentifications and
only Helicobia morionella, H. rapax, Oxysarcodexia
ventricosa and Ravinia lherminieri, all listed from
Bermuda by Williams (1958, 1959), and all very
common, widespread North American species,
seem reliable (I have seen recently collected mater-
ial of all but the latter species in USNM).
Research on the Nearctic Microcerella, evoked by
the discovery of the Bermudan species, revealed
that what has been considered a single taxon, M.
scrofa, actually covers two sibling species. Previous
illustrations of terminalia in Aldrich (1916: fig. 13),
Roback (1954: fig. 39) and Lopes (1982c: figs. 43-
47) seem to be of M. scrofa proper, while the species
figured as Hypopelta scrofa in Hallock (1940: figs.
1-2) probably is its sibling M. adelphe (the surstylus
figured by Hallock is perfectly scrofa-like, but this
may be due to the rather simple outline provided by
this author and the fact that the thornlike surstylar
process is difficult to see in strict lateral view).
Both new species of Microcerella are described
below.
Microcerella bermuda sp. n. (figs. 33-36)
Type material. — Holotype @, "Bermudas | printed] vii
8 [handwritten, black ink] J.8 [handwritten, pencil] Davis
[printed] (MCZ). Paratype, 18, "Bermuda Islands July 4
1905, T. K” [only “4” handwritten] (ZMUC). Both types
bear a label reading "Sarcophaga n. sp. det. ALDRICH”.
Description.
Male. — Head: Narrowest part of frons 0.24X
head width. Outer and inner vertical bristle well
DIL
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
developed. Two weak ocellars present, the bristles
much weaker than the pair of postocellars. Frontal
bristles (7)-8; row continuing to just above level of
apex of scape and distinctly curving laterally when
seen from above. Fronto-orbital plate otherwise
bare. Parafacial plate with a row of 4 setae in ventral
part close to eye margin. Postcranium with poster-
iormost postgenal setae and setae below occipital
foramen white; other setae black. Scape distinctly
raised above lunule, pedicel somewhat elongate,
first flagellomere 1.5X as long as pedicel. Arista
well developed, 2X as long as first flagellomere and
short plumose. Palpus black.
Thorax: Proanepisternum and postalar wall
bare. Metasternum setose. Chaetotaxy: acr = 0 + 1,
de = 4 (anteriormost 2 weak) + 3, ia = 2 (inner
posthumeral + a weak presutural) + 2, sa = 2 + 3,
pa = 2. Scutellum with 2-3 marginals (if 3 then the
two posterior ones close together), one pair of weak
discals, and one pair of apicals (slightly irregular in
the holotype).
Wing: Tegula black or brownish, basicosta yel-
low, veins yellowish brown. Costal spine about 1.5X
as long as crossvein r-m. Setal row of 14,5 just
surpassing half the distance to r-m.
Legs: Midfemoral apical pv bristles not spinelike
(no midfemoral comb), hind tibia without any elon-
gated pv or p setae.
Abdomen: Ground colour black, microtomentum
forming the usual silvery grey, tessellate and chan-
ging pattern. Tergite 1+2-3 without median mar-
ginal bristles, T4 with a weak pair, T5 with a com-
plete row of marginals. ST3-4 with long setae with
wavy or undulating tips.
Terminalia: Protandrial segment dark reddish
brown, with a few bristles on disc and a complete
row of marginals. Epandrium bright red. Cercus
and surstylus densely setose, cercus broad and plate-
like when seen in posterior (dorsal) view, S-shaped
in lateral view. ST9 (hypandrium) with a reduced
disc but with arms greatly swollen at level of gonop-
odal articulation. Gonopod shortened but rather
broad at base, partly hidden by the swollen hypan-
drial arm (lateral view), and with setae scattered
along most of the distal margin. Aedeagus with a
short basiphallus that is rigidly hinged to the dis-
tiphallus. Distiphallus with a large transversely
flattened plate carried on a narrow petiole from the
ventral surface. Juxta present, well sclerotized and
forming two arms slanting ventrally and basally.
Lateral styli tubular, median stylus weakly sclero-
tized and somewhat inflated, all styli projecting
between juxtal arms.
Length. — 10.0 mm.
Female. — Unknown.
Etymology. — A noun in apposition. Named for
the type locality.
52
Distribution. — Nearctic: Bermuda Is.
Note. — Johnson (1904) mentions a
"Sarcophaga sp. ? A smaller form of which a
number of specimens were collected by Mr. Davis,
July 8”. These specimens may or may not include
the holotype of the present species but obviously
the note refers to the very collecting trip where it
was taken (identical collector, locality and day +
month) and settles with reasonable certainty the
year when the holotype was collected as 1903, when
Mr. C. Abott Davis visited the islands.
Microcerella adelphe sp. n. (figs. 38, 40-43)
Type material. — Holotype 6, USA: Maryland, Mont-
gomery County, Rockville, 14.vi.1969, G. Steyskal
(USNM). Paratypes, Canada: Ontario, Ottawa, 1d, 4-
7.viii.1982, L. Huggert (ZML); Quebec, Mount Tremb-
lant, Lac aux Atocas, 18, 28.viii.1956, E.B. Thurman, "Bit-
ing & Sweeping” (USNM).- USA: Georgia, 18, [no date],
C. V. Riley [terminalia lost] (USNM); Minnesota, Ea-
glesnest, 18, 29.vii.1958, W. V. Balduf (USNM); New
York, Tompkins Co, [....] (illegible ?Ellio), 18,
25.viii. 1956, B. Foote, lab reared ex Polygura thyroides
(ZMUC); New York, Tompkins Co, Varna, 16,
17.vii.1946, A. Stone (USNM): North Carolina, Great
Smokie Nat. Park, Newfnd Ridge, 18, 11.vii.1941, A. L.
Melander (USNM); Virginia, Great Falls, 16, 21.vi.1931,
A. C. Melander, 19, "x.23,20”, J. M. Aldrich (both in
USNM); West Virginia, Cranberry Gla., 1, 2.vi.1955, H.
V. Weems (USNM).
Description
Male. — Very similar to M. scrofa. I have found
only few diagnostic characters, all pertaining to the
terminalia, and the description is accordingly res-
tricted to these structures. Moreover, to facilitate a
separation from M. scrofa, the description is given
as one entry of a recognition couplet. For details of
general morphology, the description of M. scrofa in
Aldrich (1916) should be consulted.
— Surstylus distally evenly rounded. Distiphallus
broadest proximal to middle (lateral view) and
with the appendages at the ventral surface
rather slender (figs 5/159 144) ae ee ee
WE RSA LAO Re En Microcerella scrofa
— Surstylus distally with a concave incision and a
thornlike process. Distiphallus broadest at or
distal to middle (lateral view) and with the
appendages at the ventral surface more com-
PACS D STORES RARI
One female has been included in the type series
of M. adelphe although I cannot separate it mor-
phologically from that of M. scrofa. Geographical
evidence, however, suggests that M. adelphe may
have a more easterly extended distribution relative
to M. scrofa (see below). A female from Ohio (Am-
herst) is accordingly judged to belong to M. scrofa.
Etymology. — A noun in apposition. From the
Greek adelphe = sister, the name refers to the
probable sister group relationship with M. scrofa.
Distribution of M. adelphe. — Nearctic: Canada
(Ontario, Quebec), USA (Georgia, Maryland, Min-
nesota, New York, Virginia, West Virginia).
Biology. — The only information available is the
breeding record from the snail Polygura thyroides
(from label of one of the paratypes from New York
State), and M. adelphe is probably predatory on
snails, as many of its congeners. Microcerella
adelphe may actually be the “undescribed species
[which] parasitizes snails” mentioned by Downes
(1965: 951). Nothing is known of the biology of M.
scrofa.
Distribution of M. scrofa. — Nearctic: Canada
(Ontario [nr. Stratford]), USA (Alabama [Bir-
mingham], Illinois [Algonquin], Indiana [Lafa-
yette], Kansas [Topeka], Louisiana [Opelousas],
Ohio [ Amherst], Texas [College Station], and Wis-
consin [state record only]). The specimens on
which this distribution is based are deposited in
USNM (US records) and ZML (Canadian record).
The apparent sympatry of M. adelphe and M.
scrofa in at least part of their range (one male of
each species from Canada: Ontario) is strong evi-
dence that the rather slight differences reflect a
reproductive barrier.
The closest relative of M. adelphe is most prob-
ably M. scrofa, as already mentioned. Both species
possess a highly modified distiphallus with lateral
sclerotized plates that extend posteriorly, thereby
forming a deep, narrow groove along the posterior
surface. Moreover, the gonopods are very character-
istic by being greatly extended anteriorly, strongly
curved upwards (or dorsally), and almost joining in
the median plane (fig. 42). The group consisting of
the sibling species adelphe + scrofa is probably the
sister group to Microcerella bermuda. The character
corroborating this hypothesis is the shape of the
gonopod. In all three species, the gonopod has
transformed from the elongate and more or less
narrow structure found in all other Microcerella
into a short or low, but very broad plate. The scat-
tered setae that cover most of the distal margin may
be used as a morphological landmark indicating the
homologue of the posterior (or dorsal) margin of
the plesiomorphic gonopod. The sister group of
adelphe + scrofa + bermuda will probably be found
among the Neotropical members of the group, as
the three remaining Nearctic species of Microce-
rella may have the Argentinean species M. muehni
PAPE: American Sarcophaginae
as their closest relative, these four species sharing
the derived black/black condition of terminalia.
Genus Spirobolomyia
Spirobolomyta Townsend, 1917a: 43. Type species: Sarco-
phaga singularis Aldrich, 1916, by original designa-
tion.
Diagnosis. — The genus Spirobolomyia may be
defined by the following character states.
1) Male mid femur apically with a row of comb-
like posteroventral spines;
2) Male abdominal sternite 5 with bristles along
the posterior margin;
3) Male abdominal sternite 5 with a pair of bristly
pads on the disc;
4) Male cercal prong bent backwards;
5) Male cercus with a sinuous lateral margin (pos-
terior view);
6) Male cercus with short spinelike setae on the
dorsal surface;
7) Basal parameral sclerite elongated (“additional
forcipes” of Lopes 1975f);
8) Aedeagus with a beaklike projection arching
over the heavily sclerotized juxta (fig. 48);
9) Aedeagal vesica beaklike and sclerotized;
10) Female abdominal sternites 6-8 fused;
11) Female abdominal tergite 6 with the median
(= dorsal) part of the posterior margin devoid
of setae and produced in a tongue-like projec-
tion (Lopes 1975f: figs. 8, 17, 26).
Townsend seldom explained the etymological
derivation of his numerous generic names, but
when he erected the genus Spirobolomyia with a
type species that had been bred from a millipede, it
seems probable that he, although he gave no men-
tion of species other than the type species, included
Sarcophaga flavipalpis as well, as this very similar
species had been "reared ... from a myriapod” (Al-
drich 1916: 258). Accordingly, he later stated that
Spirobolomyia "Ranges in two species” (Townsend
1938: 68). Downes (1965) widened the concept by
including Sarcophaga basalis and S. ohioensis in
Spirobolomyia (as subgenus of Blaesoxipha), the
biology of which is still unknown. It might be
guessed that they are all parasitoids of the millipede
genus Narceus. Note that I am here following Kee-
ton (1960), who revised the Spirobolidae and
argued that Spirobolus (consistently misspelled
Sparobolus by Aldrich 1916: 186, 258) should be
used for the group of Old World (China) species
only, while Narceus should be applied to its New
World sister group. As the name Narceus is the
oldest genus-group name within the tribe Spirobo-
lini, Narceus is the valid name in the present con-
text whether or not Keeton’s infratribal classifica-
tion is accepted.
Species of Spirobolomyia are mainly distributed
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
in the eastern and southeastern parts of the Nearc-
tic Region but extend along the east coast into the
northern part of the neotropical region. The range
extends from Canada: southern Quebec, (14 of S.
flavipalpis in ZMUC) south to Mexico: Tamaulipas
(18 of S. latissima in FSCA, see under type material
below) and Chiapas (86 29 of S. ohioensis in CNC).
The western limit of distribution seems to run
through Wisconsin, Iowa, West Virginia, and Ala-
bama (Downes 1965), and the Nearctic distribution
is closely matched by the distribution of Narceus
given in Keeton (1960).
The life habit, although only known for two spe-
cies and without any evidence that no other food
source can be utilized, is interesting insofar as very
few Diptera parasitize or prey upon millipedes.
Within the Sarcophagidae, I know of only two other
records: Specimens of Blaesoxipha beameri in
USNM (44, USA: California, Riverside) bear a
label reading "Ex Tylobolus sp. near hebes” prob-
ably referring to a species of the spirobolid genus
Hiltonius. Although this may refer to parasitism,
predation or even scavenging, the food source ne-
vertheless is a millipede. No other information is
available on the biology of B. beameri. A single male
specimen of Sarcophaga iulicida from Portugal
(BMNH), with an affiliation in the subgenus Pzer-
retia sensu stricto of Verves (1986), was bred from
a specimen of Ommatoiulus moreleti (Pape 1990).
Note that the parasitism of millipedes has not been
used as a character state corroborating the mono-
phyly of Spirobolomyia. Parasitism of a particular
taxon is difficult to handle in a character analysis as
parasites do not parasitize taxa as such, and al-
though it seems reasonable that the peculiar habit
of Spirobolomyia is evidence of phylogenetic rela-
tionship I prefer to avoid an explicit scoring.
Spirobolomyia was, like Fletcherimyia, included
in Blaesoxipha as a subgenus by Downes (1965).
However, all Spirobolomyia possess tubular styli
that are free of each other and apparently func-
tional, i.e. sperm conducting, and they do therefore
not form part of the probably monophyletic group
of species with only the median stylus tubular and
with platelike lateral styli that are fused in the
median plane. The latter group is the genus Blae-
soxipha in the sense of the present author. The
male midfemoral comb, the presence of short
spines on the dorsal surface of the male cercal
prong, the recurved male cercal prong, and the
fused female abdominal sternites 6-8, however, are
character states Spirobolomyia shares with Blae-
soxipha. At least the latter four states seem derived
with regard to the groundplan of the Sarcophagi-
nae, and including Spzrobolomyia within Blaesoxi-
pha (i.e. widening the concept of the latter) may be
theoretically sound on the present state of know-
ledge, and my reason for not doing this is, as given
for Comasarcophaga and Fletcherimyia, that the
54
monophyly of the resulting taxon seems, at least to
me, to be much more tentative than the monophyly
of the groups included. A comprehensive phyloge-
netic analysis of all sarcophagine genera is needed
to settle this issue. Lopes (1975f: 156) simply dis-
carded a relationship between Spirobolomyia and
Blaesoxipha by stating that “there is no close rela-
tionship between the two genera”, but he later
(1988a: 130) elaborated on this postulate by refer-
ring explicitly to the “long styli of glans” (= lateral
styli of the present paper) being present in species
of Spirobolomyia but absent in his Impariini (=
Blaesoxipha (in part) of the present paper). The
possession of functional lateral styli will corrobo-
rate that Spirobolomyia does not form part of the
clade here considered under the name Blaesoxipha
(unless other evidence favour a reversal), but as the
character state obviously is plesiomorphic at the
level of the Sarcophaginae, it is totally uninforma-
tive regarding the phylogenetic position of Spzro-
bolomyia relative to Blaesoxipha.
Species limits were discussed in detail by Lopes
(1975f), but the number of specimens at his dispo-
sal was rather limited and a more extensive mate-
rial has revealed another, previously undescribed
species. As only characters found in the male termi-
nalia distinguish this species from S. ohioensis, and
as species of Spirobolomyia in general are separable
only by reference to their terminalia and sternite 5,
I have restricted the description to include these
structures only. Information on general appearance
of Spirobolomyia can be found in Aldrich (1916),
Hall (1927), Townsend (1938) and Lopes (1975f).
Spirobolomyia latissima sp. n. (figs. 45-48, 50)
Type material. — Holotype d, USA: Texas, Menard,
1929, E. O. Cushing (USNM). Paratypes, USA: Texas:
168, data as holotype (14 in USNM, 2 in ZMUC); 18
v.1930, otherwise as holotype (USNM); NW Blanco Co.,
Davis Ranch, 16, 22.iv.1959, J. F McAlpine (CNC); Rea-
gan Wells, 18, 27.vi.1927, R. W. Laake (USNM); Sonora,
1d, 10. [month not interpreted].1920, O. G. Babcock
(USNM); Hidalgo Co., 78, 29.iii.-9.x.1946, USPHS dys-
entery fly trap (USNM); Hidalgo Co., 3&: 4.ii.1932, 14:
3.viii. 1934, H.J. Reinhard (3 in CNC, 1 in ZMUC); Brews-
ter Co., Big Bend National Park, Basin, 14, 14.vi.1950, R.
F. Smith (AMNH). - Mexico: Tamaulipas, 9 km W An-
tiguo-Morelos, 18, 21.vii.1954, Univ. Kansas Exp.
(ESCA).
Description
Male. — Abdomen: Sternite 5 deeply incised and
with strong, spinelike bristles along the basal (or
median) half of the posterior margin of each lobe.
A pair of raised pads with strong bristles on the
posterior, flattened surface are present on the disc.
Terminalia: Cercus very broad in posterior view,
with a sinuous outer (or lateral) margin and a
greatly swollen flaplike extension basally on the
cercal prong. A few short spines are present on the
dorsal surface and along the margin of the hump-
like lateral convexity. Cercal setae are arranged in
two groups: a smaller one close to the median line
of articulation between the cerci, and a more exten-
sive one distal to this. Aedeagus with the dorsal (or
posterior) surface raised into a narrow crest. Vesica
well sclerotized and of about the same size as the
beaklike structure arching over the juxta.
Female. — Probably very like the female of S.
ohioensis. Among nine females from Texas, Hi-
dalgo Co., 14.iii.-23.x.1946 (USNM), all caught in
dysentery fly traps, may be specimens of S. latis-
sima. | have not been able, however, to find any
differences between these and a female specimen of
ohioensis pinned with a presumed conspecific male
(USNM), and as the distribution of these two spe-
cies seems to be largely sympatric, I have not in-
cluded any females in the type series.
Etymology. — A Latin adjective, from latissimus
= the broadest. The name refers to the male cercus,
which is very broad in posterior view.
Distribution. — Nearctic: USA (Texas). Neo-
tropical: Mexico (Tamaulipas).
Females of Spirobolomyia are very similar and I
cannot at present identify female specimens to spe-
cies with any certainty. This implies a problem, as
the holotype of Sarcophaga pallipes is a female.
Aldrich (1930) examined this type and established
its relationship, but he was not able to make a
species level identification: "I can not separate [it]
from females of simgularis and several others” (p.
19). Townsend (1938) and Roback (1954) consi-
dered S. pallipes a senior synonym of S singularis,
while Downes (1965) listed pallipes as “Probably
Blaesoxipha (Spirobolomyia) sp”. Lopes (1975c)
likewise examined the type of pallipes and syn-
onymized it with Sarcophaga basalis, but his deci-
sion obviously was tentative as he "was unable to
[find] good characters to separate the female[s] of
the Spirobolomyia with red legs” (p. 548). Appar-
ently, his decision was based largely on the equal
size of the primary types, which really does not
provide especially convincing evidence as to their
conspecificity. When Lopes (1975f) later produced
a key to species of Spirobolomyia, he did not include
the females and he did not examine any female
specimens of S. ohioensis. For the present paper, I
have accepted the synonymy of S. basalis with S.
pallipes, awaiting more information on diagnostic
female character states.
Roback (1954) illustrated the aedeagus and ab-
dominal sternite 5 of S. singularis (as pallipes), S.
flavipalpis (as flavipes, error) and S. basalis. Lopes
(1975f) gave a detailed description of these species
PAPE: American Sarcophaginae
as well as of S. ohioensis and provided illustrations
of various parts of both male and female terminalia.
Both authors should be consulted for additional
information. The key of Lopes (1975f), however, is
not easy to use and the posterior view of the male
cerci is provided for two species only (that of S.
basalis even at a slightly skewed angle). The poste-
rior outline of the male cerci provides a very easy
means of identification and has the advantage that
this attribute often is visible even in specimens with
only partly spread terminalia. For the present
paper, I have provided illustrations of the posterior
cercal outline for all species (figs. 49-53), and for
completeness a couplet is provided to separate S.
latissima from the very similar S. ohioensis:
— Aedeagus with long vesica and a crest running
along the dorsal surface (fig. 48). Cercus very
broad in posterior view (figs. 46, 50). Setae of
cercal base separated into two groups by an area
with no or very few setae: A small group close
to the suture between the cerci and a larger
group along the outer (or lateral) margin (fig.
AO sand eds bldg net se latissima
— Aedeagus with shorter vesica (Lopes 1975f: fig.
35), dorsal surface distinctly keeled but not
raised into a crest. Cercus narrower in posterior
view (fig. 49). Setose part of cercal base without
any interruption, setae forming a continuous
cover from suture between the cerci and out
along the outer (or lateral) margin..........
Genus Tripanurga
Tripanurga Brauer & Bergenstamm, 1891: 367. Type spe-
cies: Sarcophaga albicans Wiedemann, 1830, by desig-
nation of Townsend (1916: 9).
Thelylepticocnema Townsend, 1917a: 43. Type species:
Sarcophaga incurva Aldrich, 1916, by original designa-
tion.
Glaucosarcophaga Townsend, 1917a: 45. Type species:
Glaucosarcophaga knabi Townsend, 1917a [= Sarco-
phaga albicans Wiedemann, 1830], by original desig-
nation.
Metoposarcophaga Townsend, 1917a: 46. Type species:
Sarcophaga pachyprocta Parker, 1916 [junior primary
homonym of pachyprocta Hagen, 1881; = a species of
the Sarcophaga importuna Walker complex], by orig-
inal designation. Syn. n.
Zygastropyga Townsend, 1917b: 191. Type species: Zy-
gastropyga aurea Townsend, 1917b, by original desig-
nation.
Cacotrophus Reinhard, 1947: 99. Type species: Cacotro-
phus beameri Reinhard, 1947, by original designation.
Erucophaga Reinhard, 1963a: 75. Type species: Eruco-
phaga triloris Reinhard, 1963a, by original designa-
tion. Syn. n.
Diagnosis. — The genus Tripanurga may be de-
fined by the following character states:
1) Aedeagal basiphallus with an epiphallus-like
process;
2) Aedeagal basiphallus shaped as an elongated
55
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
and narrow tube, strongly contrasting to the
compact distiphallus (T. aurea showing a se-
condary thickening);
3) Aedeagal distiphallus compact and more or
less globular;
4) Aedeagal distiphallus with a fringe of filiform
processes at the anterior margin (Lopes 1978b:
figs. 6-8; the present figs. 57, 61);
5) Parameral bristle slightly flattened;
6) Ejaculatory apodeme large;
7) Male cercal prong bent backwards.
The groundplan of the probably monophyletic
group Paramacronychiinae + Sarcophaginae is
characterized by an almost completely reduced epi-
phallus, but in all species of Tripanurga the prox-
imal part of the basiphallus is extended into a dis-
tinct process that is broad at the base, tapers
gradually and slants backwards or posteriorly (Ro-
back 1954: figs. 316-335; Lopes 1978b: fig. 6; She-
well 1987: fig. 58c; the present figs. 57, 61). Shewell
(1987) applied the term epiphallus to this structure,
but I prefer to avoid it as the basiphallic process in
Tripanurga cannot be considered homologous to
the epiphallus of the Miltogrammatinae and Calli-
phoridae but rather to its base.
Character states 3-5 are here considered autapo-
morphic, and probably state 2 as well as the condi-
tion in T. aurea, which possesses a rather strong
basiphallic tube (fig. 61), is judged to be secondary.
The shape of the distiphallus may indicate a trans-
formation with a gradual increase in the size of the
dorsal humps: importuna-type — villipes-type >
sulculata-type — aurea-type (compare Roback
1954: plates 23-24, and the present figs. 57, 61).
The unique (i.e. autapomorphic) shape of the au-
rea-type distiphallus makes a reversed polarity
highly improbable. If this is accepted, T. aurea can-
not be considered the sister group of all other Tri-
panurga and the broad basiphallus found in this
species has to be considered a secondary specializa-
tion (i.e. apomorphic).
The genus Tripanurga was described by Brauer &
Bergenstamm (1891) to contain "1. albicans Wd.
(Sarcophaga). 2. dimidiata Wd. 3. bicolor S.M.C.
Bras” (p. 367). The latter name is apparently a
nomen nudum while the second probably is Wiede-
mann’s (1830) Sarcophaga dimidiata, which cur-
rently is regarded as a species of Notochaeta, e.g. by
Lopes (1969a). Two years later Brauer & Bergen-
stamm (1893) included (or mentioned) albicans
only, and Townsend (1916) designated this as the
type species, probably without having seen the type
or any conspecific specimens. Townsend spent six
months of 1928 visiting European and American
museums and examining a large number of types,
and he later published the results in various papers.
For Sarcophaga albicans, he gave the type deposit-
ories as: “Male Ht in Wien and male Pt in Lima”
56
(Townsend 1931: 75), which would have been an
effective lectotype designation if only a single syn-
typic specimen had been present in Vienna
(NMW). Townsend probably incorporated one of
the original type specimens in his own collection,
and when he later moved from Lima to Washing-
ton, D.C., bringing his collection along, the para-
type (which with the present lectotype designation
automatically becomes a paralectotype) was depos-
ited in the USNM (Townsend 1937b: 221). Aldrich
went to Europe in 1929, i.e. after Townsend, but he
published his results a year earlier, although with-
out designating any of the specimens examined as
lectotypes. He did, however, give an extensive des-
cription of two males of Sarcophaga albicans in
NMW, "Undoubted types” (Aldrich 1930: 8). As
both these males are still present in NMW, and as
Townsend’s visit in Vienna antedated that of Al-
drich, Townsend must have incorporated one of
three original syntypic males in his own collection
and not one of the two seen by Aldrich as stated by
Lopes (1978b). One of these two remaining speci-
mens was then chosen by Townsend to be the hol-
otype. This is to be considered as a valid lectotype
designation only if the specimen in case can be
properly recognized, but as Townsend neither la-
belled one of the males in question (he never la-
belled his ‘designated’ lectotypes) nor cited any in-
formation that could refer to one particular
specimen, I have simply regarded Townsend’s de-
signation as invalid and chosen one of the two
available specimens, which I have labelled as lecto-
type. This male is in good condition except that it
lacks right first flagellomere + arista, left foretar-
sus, right midleg, left hind leg, right hind tarsus,
and has the terminalia dissected and glued to the
lowermost label. It is labelled Brasilien” (printed)
and “albicans Wd J 48 Coll. Winthem” (last two
words printed), and its identity as Tripanurga albi-
cans is herewith verified. The two paralectotypes in
USNM and NMW respectively have been labelled
as such.
Silvestri (1903) described the species Tripanurga
termitophila, probably on a single female which he
recovered and apparently bred from a termite nest.
The holotype has never been revised [I have not
tried to locate this specimen, which is from Santa
Ana, Misiones, in Tucumàn Province, Argentina,
and not just “America Meridionalis” as stated by
Lopes (1969a: 47)] and Tripanurga has in practice
remained monotypic since Brauer & Bergenstamm
(1893) excluded (or omitted) Sarcophaga dimidiata.
Some authors, however, have stressed the strong
similarity to Metoposarcophaga and related taxa.
Aldrich (1930) mentioned that Sarcophaga albicans
resembled the type species of both Metoposarco-
phaga and Zygastropyga, and Roback (1954: 81)
mentioned that Tripanurga "will probably fall
within the genus Metoposarcophaga’ as T. albicans
seems to be “very close to the subgenus Zygastro-
pyga”. Neither author, however, made an explicit
synonymization. Downes (1965) employed a broad
concept of Metoposarcophaga, although he did not
include Erucophaga, but he probably did not con-
sider Tripanurga at all as T. albicans has not yet
been found north of Mexico. Lopes (1969a) listed
Sabinata, Tripanurga and Zygastropyga as distinct
genera, while Shewell (1987) separated both Thely-
lepticocnema and Erucophaga from Metoposarco-
phaga. I think that using Tripanurga in the more
inclusive sense presented above should be preferred
for the following two reasons: Monotypic genera as
Tripanurga (with T. albicans only) and Erucophaga
are avoided, and the monophyly of the resulting
taxon is better corroborated. Accepting a mono-
typic Tripanurga will leave Metoposarcophaga, e.g.
sensu Downes (1965), without any assumed apo-
morphies not shared by T. albicans, and further
splitting with the creation of additional monotypic
genera has to be accepted. The genus Erucophaga
provides a somewhat different case as the straight
or slightly anteriorly curved male cercus of tri/orzs
is unique within Tripanurga (in the broad defini-
tion) and may be plesiomorphic relative to the
remaining species where the cercus is distinctly
curved backwards. This would be an argument for
treating this species as the sister group to all other
Tripanurga, and it may, therefore, be an argument
for maintaining the genus Erzcophaga. I still think,
however, that this is unwarranted simply because
the little new information presented (= the possi-
ble monophyly of Tripanurga excl. triloris) is made
up for by a similar loss (= the monophyly of Tri-
panurga incl. triloris). Moreover, as no well corrob-
orated sister group of Tripanurga incl. triloris has
been proposed, the monophyly of Tripanurga excl.
of triloris will be highly tentative, and one ends
with nothing but another genus-group name to
handle. Actually, if Argoravinia turns out to be the
best candidate for the sister group, as discussed
below, the cercal condition in treloris must be con-
sidered as derived from the bent cercus of its con-
geners.
In conclusion, the monophyly of Tripanurga in
the present, broadened sense is corroborated by
some very convincing autapomorphic character
states of the male copulatory apparatus, which
furthermore makes the taxon easily recognizable, at
least in the male sex.
Some of the character states listed in the diagno-
sis may be apomorphic at a higher (i.e. more inclu-
sive) level. The large ejaculatory apodeme, for ex-
ample (Hallock 1940: fig. 13; Lopes 1978b: fig. 6;
Shewell 1987: fig. 58c), is likewise found in the
genus Argoravinia (although not quite as large) and
it may be taken as evidence for considering these
two genera as sister groups. Hall (1933: 255) even
stated that Argoravinia “appears greatly like the
PAPE: American Sarcophaginae
genus Metoposarcophaga”. Moreover, a rather
broad male front (although with proclinate orbital
bristles in albicans and triloris only) is likewise
shared with Argoravinia, and in all species of Tri-
panurga except T. triloris, the male cercus is bent
backwards or posteriorly (figs. 54, 58), somewhat
like the condition found in Argoravinia. The almost
straight male cercus in T. triloris may then repres-
ent a derived state.
Tripanurga guatemala sp. n. (figs. 54-57)
Type material. — Holotype 8, Guatemala: Antigua, El
Salto, 1.v.[no year], J. M. Aldrich (USNM). Paratypes,
Guatemala: Antigua, 14, 14.v.1923, E.G. Smyth (ZMUC);
Guatemala City, 18, vii.1923, E. G. Smyth (USNM).
Description
Male. — Head: Narrowest part of frons 0.27-
0.29X head width. Outer and inner vertical bristle
well developed, fronto-orbital plate densely setose
and without proclinate orbitals. Uppermost frontal
bristle strong and reclinate. Upper half of parafacial
plate with some scattered setulae, lower part with
a row of setulae close to eye margin. All postcranial
setae, including those on hypostomal bridge, black.
Head entirely silvery microtomentose. Antennal
pedicel blackish with brownish base and apical mar-
gin, first flagellomere blackish grey. Palpus black.
Wing: Tegula black, basicosta yellow. Vein r,
setose from base almost to the knob at level of
subcostal bend/costal break, r,,; setose from base
almost to crossvein r-m.
Legs: All femora with long setae and without av
or pv bristles. All tibiae, especially mid- and hind-
with dense villosity.
Abdomen: Tergite 1+2-3 without median margi-
nal bristles, T4 with a median pair, and T5 with a
complete row of marginals.
Terminalia: Colour red. Cercal prong slender,
prongs closely adjoining in the distal half leaving a
small window at base. Surstylus broad, with a fringe
of long flattened setae.
Length. — 11.0-13.0 mm.
Female. — Unknown.
Etymology. — A noun in apposition. Named for
the type locality.
Remarks. — T. guatemala is most probably the
sister species of T. villipes, the monophyly of this
group being well corroborated by at least three
distinctive character states that may be derived rel-
ative to the groundplan of Tripanurga:
1) Postcranial setae all black;
2) Alllegs equipped with dense, long pilosity and
no anteroventral or posteroventral bristles;
3) Strongly convex ventral margin of the sursty-
DI
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
lus with a row of closely set, strong, and elon-
gated setae.
Tripanurga aurea (figs. 58-66)
Distribution. — Nearctic: USA (Arizona, Cali-
fornta), Mexico (Baja California, Sonora). Neotrop-
ical: Mexico (Guerrero, Jalisco, Morelos), Nicara-
gua.
The reason for discussing the present species is
that it seems to be split into two populations, a Baja
Californian population and a mainland population,
with some striking morphological differences be-
tween the two. Indeed, these differences were
thought to indicate a specific separation until I
recovered specimens from northern Baja California
showing intermediate character states and reticu-
late character state combinations. Two characters
showing especially pronounced differences, the out-
line of the surstylus and the shape of the male
abdominal sternite 5, are described in more detail
below.
The typical Baja Californian form (figs. 58-63)
has a slight posteroventral concavity of the sursty-
lar margin (best seen in lateral view as in fig. 58),
and the lobes of the male abdominal sternite 5 are
strongly swollen apically and transversely flattened
posteriorly (fig. 62). Specimens of the mainland
form have the surstylus deeply incised from the
posteroventral margin (figs. 64, 65), and the lobes
of abdominal sternite 5 are flat and evenly rounded
(fig. 66), i.e. they represent unmodified lobate. ex-
tensions of the sternal disc. The aedeagus is com-
pletely identical in both populations (fig. 61), and
no differences have been found in the female sex.
It is interesting that the Baja Californian form
does not seem to be just a peripheral isolate that has
diverged from its plesiomorphic ancestral popula-
tion on the mainland. Both populations (i.e. gene
pools) have probably diverged from their common
(hypothetical) ancestor by the acquisition of at least
one autapomorphic character state, the Baja Cali-
fornian form having developed the obviously apo-
morphic swollen lobes of abdominal sternite 5 and
the mainland form the incised surstylus. The two
forms are morphologically homogeneous through-
out their range, but in the area where they occur
simultaneously (northern Baja California), inter-
mediary specimens and specimens with transposed
character state combinations, i.e. incised surstylus
and swollen lobes of ST5, may be caught at the same
locality. Therefore, reproductive isolation does not
seem to have been established, and rather than
proposing a formal subspecific name for the Baja
Californian population, I prefer to use the term
“form”, awaiting further studies of the popula-
tion(s) showing intermediary conditions.
58
Material examined.
Type material. — USA: Holotype 8, Arizona, Tempe,
[no date], Webster (USNM).
Other material. — Mexico: Sonora, Alamos, 4d, 20-
27.11.1963, P. H. Arnaud, Jr. (USNM); Jalisco, Labarca, 14
19, 3.x.1934 [no collector] (UNAM); Guerrero, Arcelia,
16, 21.x.1947, W. G. Downs (UNAM); Morelos, 5 km
SW Amacuzac, 36, 8.vi.1979, J. Butze (2 in UNAM, 1 in
ZMUC); Baja California Norte, Arr. Santo Domingo, 5.7
mi E Hamilton Ranch, dam site, 428 529, 23.iv.1963, H.
B. Leech & P. H. Arnaud, Jr. (FSCA); Golfo California, Isla
Angel de la Guarda, Puerto Refugio, 24, 6.11.1986, F. Arias
(UNAM, ZMUC); Golfo California, Isla Espiritu Santo,
18 19, 11.1.1987, L. Cervantes (UNAM). - Nicaragua:
Chinandega, 18, [no date], Baker (USNM).
ACKNOWLEDGEMENTS
I am most grateful to N. E. Woodley (USNM) for
facilitating my access to the collections during a
one-year fellowship at the Smithsonian Institution
and for supplying important information on Ber-
mudan Sarcophagidae. I thank W. L. Downes, Jr.,
Lansing, Michigan, for sharing much unpublished
information on Nearctic Sarcophagidae, and K.
Rognes, Hafrsfjord, Norway, for a sober and very
professional critique. Thanks also to W. N. Mathis,
F.C. Thompson, and N. E. Woodley (all USNM) for
heuristic discussions of various aspects of generic
definitions. I acknowledge the time spent by D.
Grimaldi (AMNH), G. E. Shewell and B. E. Cooper
(CNC), C. Vogt (MCZ) and H. V. Weems, Jr.
(FSCA) who kindly introduced me to the collections
of their respective institutions during short visits
and made prompt loan arrangements. Other loans
were kindly supplied from L. Fernandez (UNAM),
A. O. Bachmann (MACN), and R. Danielsson
(ZML). Mary E. Petersen (ZMUC) made valuable
linguistic corrections.
Grants from the Smithsonian Institution, the
Danish Natural Science Research Council (grant
no. 11-6152), and the CanaColl Foundation are
gratefully acknowledged.
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PAPE: American Sarcophaginae
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59
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
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Lopes, H. S., 1982c. On some Microcerellini (Diptera,
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PAPE: American Sarcophaginae
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Received: 15 November 1989
Revised version accepted: 12 April 1990
61
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 1-4. Comasarcophaga spp. Aedeagus plus right paramere + gonopod, lateral view. 1, C. longespinus. 2, C. prolepsis.
3, C. texana. 4, C. nexilis. Scale: 0.1 mm.
62
PAPE: American Sarcophaginae
E
10 N.
11
Figs. 5-12. Comasarcophaga spp. Male cerci. Figs. 5-8, right cercus, lateral view. Figs. 9-12, cerci, posterior view. 5, 9,
C. longespinus. 6, 10, C. prolepsis. 7, 11, C. texana. 8, 12, C. nexilis. Scale: 0.1 mm.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
55
GT LLC
mn,
mi
5:
nt mme — N
(a x
<
Ò) i
ff
1m i DI
Pd, at m
rm
In
Ama,
MON MIT
Figs. 13-18. Fletcherimyia spp. Figs. 13-17, aedeagus plus right paramere + gonopod, lateral view. Fig. 18, distiphallus,
ventral view. 13, F. abdita. 14, F. celarata. 15, F. fletcheri. 16, F. jonesi. 17, 18, F. rileyi. Abbreviations: J = juxta, LS =
lateral styli, MS = median stylus, V = vesica. Scale: 0.2 mm.
64
PAPE: American Sarcophaginae
Figs. 19-23. Fletcherimyia spp. Male abdominal sternite 5, as seen from below between sternite 4 and margins of tergite
5. 19, F. abdita. 20, F. celarata. 21, F. fletcheri. 22, F jonesi. 23, F. rileyi. Scale: 0.4 mm.
65
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
|
j
Figs. 24-28. Fletcherimyia spp. Outline of male cerci + right surstylus, posterior view. Dotted line on right cercus delimit
area devoid of setae. 24, FE abdita. 25, F. jonesi. 26, F celarata. 27, F. fletcheri. 28, F. rileyi. Scale: 0.2 mm.
66
PAPE: American Sarcophaginae
Figs. 29-32. Microcerella acridiorum. Male terminalia. 29, Aedeagus, lateral view, 30, Right paramere + gonopod, lateral
view, 31, Outline of cerci. posterior view. 32, Outline of right cercus + surstylus, lateral view. Scale: 0.2 mm.
67
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 33-36. Microcerella bermuda. Male terminalia. 33, Right cercus + surstylus, lateral view. 34, Outline of cerci,
posterior view. 35, Right paramere + gonopod, lateral view, plus posterior end of right hypandrial arm (hatched). 36,
Aedeagus, lateral view. Abbreviations: J = juxta, LS = lateral styli, MS = median stylus. Scale: 0.1 mm.
68
PAPE: American Sarcophaginae
Figs. 37-44. Microcerella spp. Male terminalia. Figs. 37, 39, 44, M. scrofa. Figs. 38, 40-43, M. adelphe. 37, right cercus
+ surstylus, lateral view. 38, right cercus + surstylus, lateral view. 39, right cercus (in part) + surstylus, posterior view.
40, right cercus (in part) + surstylus, posterior view. 41, paramere. 42, right gonopod (stippled) and posterior part of
hypandrial arm, lateral view. Fig. 43, aedeagus. Fig. 44, distiphallus. Scale: 0.1 mm.
69
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990 ,
Figs. 45-48. Spirobolomyia Latissima, male terminalia. 45, Abdominal sternite 5, ventral view. 46, Cerci, posterior view,
setae of left cercus omitted. 47, Right paramere + gonopod, lateral view. 48, Aedeagus. Abbreviations: B = basal
parameral sclerite, G = gonopod, J = juxta, MS = median stylus, P = paramere, V = vesica. Scale: 0.2 mm.
70
PAPE: American Sarcophaginae
50
Figs. 49-53. Spirobolomyia spp., outline of male cerci, posterior view. 49, S. ohioensis. 50, S. latissima. 51, S. basalis. 52,
S. singularis. 53, S. flavipalpis. Scale: 0.2 mm.
71
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 54-57. Tripanurga guatemala, male terminalia. 54, Right cercus + surstylus. 55, Outline of cerci. posterior view.
56, Right paramere + gonopod, lateral view. 57, Aedeagus. Scale: 0.1 mm.
72
PAPE: American Sarcophaginae
Figs. 58-61. Tripanurga aurea, male terminalia. 58, Left cercus + surstylus, lateral view. 59, Right paramere + gonopod,
lateral view. 60, Outline of cerci, posterior view. 61, Aedeagus, lateral view. Scale: 0.1 mm.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 62-66. Tripanurga aurea, Details of male terminalia. Figs. 62, 63, Baja Californian form. Figs. 64-66, mainland form.
62, Lobes of abdominal sternite 5, posterior view. 63, Right surstylus, posterior view. 64, Distal part of right surstylus,
posterior view. 65, Outline of left surstylus, lateral view. 66, Lobes of abdominal sternite 5, posterior view. Scale: 0.1
mm.
74
VIRGINIJUS SRUOGA
Zoologijos katedra VPI, Vilnius
SEVEN NEW SPECIES OF ELACHISTIDAE
(LEPIDOPTERA) FROM THE USSR
Sruoga, V., 1990. Seven new species of Elachistidae (Lepidoptera) from the USSR. Tijd-
schrift voor Entomologie 133: 75-84, figs. 1-26. [ISSN 0040-7496]. Published 31 July 1990.
Seven new species of Elachistidae are described from Tadzhikistan and Turkmeniya (Soviet
Central Asia) and Primorskiy Kray (Soviet Far East). They belong to the genera Perittia
Stainton, Elachista Treitschke, Biselachista Traugott-Olsen & Nielsen and Cosmiotes Cle-
mens. The male external features and genitalia are described and figured in detail. The
distribution is mapped. A historical review of studies on USSR Elachistidae is provided in
the introduction.
V. Sruoga, Zoologijos katedra VPI, g. Studentu 39, Vilnius 34, Lithuania.
Keywords. — Elachistidae, new species, USSR, Central Asia, Primorskiy Kray.
INTRODUCTION
Elachistidae larvae are typically leafminers of
herbaceous, monocotyledonous plants, belonging to
the families Poaceae, Cyperaceae and Juncaceae.
Some species are leafminers of herbaceous dicoty-
ledons belonging to the Boraginaceae, Lamiaceae,
Asteraceae and Cistaceae (Traugott-Olsen &
Nielsen 1977). Only in a few cases Elachistidae
larvae mine leaves of woody Caprifoliaceae
(Lonicera). Elachistidae larvae form galleries or
blotches between the upper and lower epidermis
layers and feed on the green parenchyma. The adult
moths are small, with a wingspan of 6 to 13 mm.
The forewings of the majority of species are brown-
ish-grey with one fascia or two fasciae. Some Ela-
chistidae are unicolorous brown, white or cream.
The family is widely distributed. However, revi-
sions of Elachistidae are only published for the
Nearctic fauna (Braun 1948) and the North Euro-
pean fauna (Traugott-Olsen & Nielsen 1977).
The last decade a considerable number of taxo-
nomic papers has been published with descriptions
of new species from western and central Europe,
northern Africa, the Canary Islands, western and
central Asia (Parenti 1978, 1981, Traugott-Olsen
1985a, 1985b, 1988, Nielsen & Traugott-Oben
1978b, 1981, 1987, Whitebread 1984) and also from
Japan (Kuroko 1982, Parenti 1983). The Elachisti-
dae fauna of the USSR has been poorly studied. In
the Keys to the insects of the European part of the
USSR” (Falkovitsh 1981) 77 species of Elachistidae
have been treated, including figures of their geni-
talia. This key was entirely based on Traugott-Olsen
& Nielsen’s book (1977) on the Fennoscandian and
Danish fauna, and did not provide original data on
Elachistidae from the USSR territories.
As early as the mid-19th century, five species of
Elachistidae were recorded from Latvia, i.e. Elachista
quadripunctella (Hübner), E. luticomella Zeller, E.
pollinariella Zeller, E. albifrontella (Hübner) and E.
argentella (Clerck) (Lienig 1846). Nine other spe-
cies were added by Nolcken (1871), te. Elachista
regificella Sitcom, E. humilis Zeller, E. pulchella
(Haworth), E. bisulcella (Duponchel), E. cerusella
(Hübner), E. gleichenella (Fabricius), Biselachista
serricornis (Stainton), B. wtonella (Frey) and B.
albidella (Nylander). Nolcken (1871) also recorded
16 species for the Estonian fauna. Twenty-four spe-
cies were included in the Catalogue of Russian Le-
pidoptera by Ershov & Fild (1870). Teich (1889,
1893, 1899) recorded five additional species of Ela-
chistidae for the Baltic fauna: Elachista eleochariella
Stainton, E. pullicomella Zeller, E. biatomella Stain-
ton, E. dispilella Zeller and Cosmiotes freyerella
(Hiibner). The Estonian entomologist Petersen
(1924) mentioned 25 species for the Estonian fauna,
four of which (Elachista apicipunctella Stainton, E.
megerlella Stainton, E. poae Stainton and E. pome-
rana Frey) were recorded for the first time for the
eastern baltic region (the present Baltic republics).
Some additional data on the Baltic Elachistidae were
provided by Saar (1930), Palionis (1932), Brandt
(1942), Prüffer (1947) and Kuusik (1962). Elachis-
tidae from other eastern European regions are dealt
with by Schille (1931): he noted 32 species from
eastern Poland (now parts of the Ukraine).
In recent years investigations on Elachistidae of
various regions in the USSR (including the Baltic
republics) have been intensified. Sulcs & Sulcs
(1983, 1984, 1987), Savenkov (1984, 1987) and
Ivinskis et al. (1985) recorded 48 species of Elachis-
tidae for the three Baltic republics, and two more
75
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
species will be added shortly (N. Savenkov pers.
comm.). From other regions of the European part
of the USSR the studies in the Vyborg reserve near
Leningrad are worth mentioning: 17 Elachistidae
were recorded here (Sukhareva & Falkovitsh 1984).
Further, Sinev (1988) found two species, Elachista
nielswolffi Svensson and Cosmuotes exactella Her-
rich-Schaffer, in the Murmansk region in 1979-
1980, amongst a considerable number of Lepidop-
tera species. Material of the Crimean Elachistidae is
still under study (Yu. Budashkin pers. comm.).
Hitherto, only one work dealt with the Elachis-
tidae of the Asiatic part of the USSR (Falkovitsh
1986), in which one new genus (Kuma Falkovitsh)
and three new species (Kumia integra, Elachista
ilicrina and E. manca) were described.
From this review it is clear that the Elachistidae
fauna of the USSR is very imperfectly known, and
further studies are urgently needed for a better
understanding of this rich and interesting fauna.
This paper contributes with the description of
seven new species from two regions in the Asiatic
part of the USSR.
MATERIAL AND METHODS
Methods for preparation of genitalia largely fol-
low Falkovitsh & Stekolnikov (1978). The genitalia
of the Cosmiotes species are figured here with the
valvae in situ, because of difficulties in spreading
them without damage. The genitalia were studied
with a Biolam microscope and a MBS-10 stereo-
microscope, using the drawing apparatus according
to Gorodkov (1961) for the line figures.
The material studied has been collected during
expeditions to largely unexplored and little known
areas of Soviet Central Asia. In addition material
from Tadzhikistan and Primorskiy Kray collected
by R. Puplesis in 1982 to 1986 has been studied.
Locality names are spelled in accordance with the
Times Atlas of the World (comprehensive edn.
1975 and later).
Fig. 1. Perittia biloba, holotype: male genitalia. Scale 0.1
mm.
76
The type specimens are deposited in the collec-
tion of the "Minology” working group, Department
of Zoology, Pedagogical Institute, Vilnius, Lithua-
nia (MG) and in the collections of the Zoological
Institute of the USSR Academy of Sciences in Le-
ningrad (ZIAS).
TAXONOMY
Perittia biloba sp. n. (figs. 1, 25)
Type material. — Holotype g: USSR, Tadzhikistan, 30
km N Dushanbe, env. Varzob (Kondara), 27.vi.1986, R.
Puplesis (MG).
Diagnosis. — Related to P. lonicerae (Zimmer-
man & Bradley), distinguished from this species by
the shape of the valvae and juxta, the basal part of
the aedeagus and the shape of the tegumen.
External characters. — Male (female unknown).
Forewing length 3.0 mm. Frons, neck tufts and
scape white, some brown scales. Antenna white and
brown ringed. Labial palp slightly mottled by white
and brownish scales. Thorax and tegulae strongly
mottled by white and brown scales. Forewing,
ground colour white, strongly mottled due to brown
tipped scales; basal part of costa densely covered by
dark brown scales; cilia grey; cilia line indistinct.
Hindwing brownish; cilia light creamy grey.
Male genitalia (fig. 1). — Uncus lobes small, with
short setae. Gnathos more or less rounded when
ventrally viewed. Tegumen widening basally. Valva
short and broad, costa strongly sclerotized, narrow,
convex, slightly setose, sacculus curved to angular,
cucullus distinct but narrow, setose. Juxta U-shaped,
lobes long, broad, distally with acute spine-like pro-
cess; digitate process very small, slightly setose.
Vinculum typical for the genus, weakly developed.
Aedeagus strongly sclerotized, slightly tapering to
distal end, two triangular lobes at base.
Biology. Unknown. Holotype caught in June.
Distribution (fig. 25). — Only known from the
holotype, Tadzhikistan.
Elachista fuscofrontella sp. n. (figs. 2-4, 26)
Type material. — Holotype @: USSR, Primorskiy Kray,
10 km S Slavyanka (Ryazanovka), 5.viii.1983, R. Puplesis
(MG).
Diagnosis. — Closely related to E. albifrontella
(Hiibner). Differs in having a darker frons and
labial palp, aedeagus curved in basal part without
cornuti, saccus gradually tapering, digitate process
club-shaped, widest above the middle, juxta lobes
not dilate.
External characters. — Male (female unknown).
Forewing length 3.0 mm. Frons, neck tuft greyish
brown, silvery lustre. Antenna greyish and dark
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Figs. 2-4. Elachista fuscofrontella, holotype. 2-3, male genitalia ventral view; 4, male genitalia lateral view. Scale 0.1 mm.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 5-8. Elachista multipunctata, holotype, male genitalia. 5-6, ventral view; 7, lateral view; 8, aedoeagus. Scale 0.1 mm.
78
brown ringed, slightly serrate distally. Labial palp
whitish from above, dark brown from below.
Thorax and tegulae greyish brown, some silvery
lustre. Forewing, ground colour dark grey, strongly
mottled, scale tips almost black; medially inter-
rupted, silvery, curved fascia before middle above
fold; tornal and costal spots whitish, apical spot,
shiny, silvery, not regular in shape; cilia brownish.
Hindwing brown; cilia brownish.
Male genitalia (figs. 2-4). — Uncus deeply in-
dented, lobes large, setose. Gnathos rounded; costa
of valva strongly sclerotized, short prominent
hump. Juxta, lobes not apically dilate; digitate pro-
cess club-shaped, very wide above middle. Vincu-
lum, not long, without medial ridge; saccus gradu-
ally narrowing. Aedeagus curved in basal part,
gradually tapering, no cornuti.
Biology. — Unknown. Holotype caught in Au-
gust.
Distribution (fig. 26). — Primorskiy Kray.
Elachista multipunctata sp. n. (figs. 5-8, 25)
Type material. — Holotype @: USSR, Tadzhikistan, 30
km N Dushanbe, env. Varzob (Kondara), 20.viii.1986, R.
Puplesis (MG).
Diagnosis. — Very closely related to E. maculata
Parenti. Differs in having the juxta lobes and dig-
itate process widening and the aedeagus with a
large cornutus. Easily distinguished from related E.
pollinariella Zeller by shape of the uncus and juxta.
External characters. — Male (female unknown).
Forewing length 3.4 mm. Frons, neck tufts white.
Antenna whitish and creamy ringed, pecten white.
Labial palp white from above, brownish from
below. Thorax white; tegulae white, some brown
scales. Forewings, ground colour white, mottled due
to brown tipped scales, basal part of costa covered
by dark brown scales; cilia white, distal part grey;
cilia line more or less distinct by dark brown tipped
scales. Hindwing brownish white; cilia greyish
white.
Male genitalia (figs. 5-8). — Uncus deeply in-
dented, lobes strongly curved towards valvae. Gna-
thos oval. Valva widest in the middle, costa with
convex lobe in the middle. Juxta, lobes wide, apically
tapering; digitate process short, wide, slightly se-
tose. Vinculum short; saccus strongly curved. Aedea-
gus strongly sclerotized, stout, sclerotized humps
past the middle and apical, one large cornutus.
Biology. — Unknown. Holotype caught in Au-
gust.
Distribution (fig. 26). — Tadzhikistan.
Elachista megagnathos sp. n. (figs. 9-12, 26)
Type material. — Holotype 4: USSR, Primorskiy Kray,
20 km E Ussuriysk (Gornotayeznoe), 6.vii.1982, R. Puple-
SRUOGA: New species of Elachistidae
sis (MG). Paratype: 1 4, same data, 7.vii.1982 (ZIAS).
Diagnosis. — Closely related to E. gleichenella
(Fabricius), easily distinguished from this species
by the very large gnathos, shape of the valva and the
basal extensions of the aedeagus. Uncus in contrast
to E. gleichenella deeply indented, digitate process
narrower.
External characters. — Male (female unknown).
Forewing length 2.6 mm (holotype), 2.9 mm (par-
atype). Head grey-brown, shining silvery. Antenna
brownish to brown, distal half slightly serrate. La-
bial palp descending, whitish, shining from above,
brownish from below. Thorax grey-brown, shining
silvery. Forewing, ground colour blackish brown
with bronze lustre, silvery marks with goldish lus-
tre, basal spot reaching costa, medial fascia distinct,
slightly widened in anal end, distal part of forewing
with second, V-shaped fascia, cilia greyish brown.
Hindwing brown; cilia greyish brown.
Male genitalia (figs. 9-12). — Uncus deeply in-
dented, lobes with small setae. Gnathos prominent,
very large. Valva widest in middle, costa convex,
cucullus setose. Juxta, apical part of lobes narrow-
ing, triangular; digitate process typical for majority
of the genus, broad, short with fine setae. Vinculum
not long, more or less triangular. Aedeagus strongly
sclerotized, characteristically bent just below distal
end, basally extended into two lateral arms.
Biology. — Unknown. Specimens were caught in
July.
Distribution (fig. 26). — Primorskiy Kray.
Biselachista spinigera sp. n. (figs. 13-16, 25)
Type material. — Holotype g: USSR, Turkmeniya,
western Kopet Dag, 30 km E Kara-Kala (Juvan-Kala),
18.viii.1988, V. Sruoga (MG).
Diagnosis. — Closest related to B. serricornis
(Stainton). Distinguished by the long and distinct
distal spine of the sacculus and the shape of the
valvae and gnathos.
External characters. — Male (female unknown).
Forewing length 3.7 mm. Frons and neck tufts
white. Antenna brownish-creamy, distal part of dis-
tal half slightly serrate, pecten white. Labial palp
white, basal part brownish from below. Thorax
white; tegulae white, some brownish scales. Fore-
wing, ground colour white, three brown spots, one
behind the middle, near costal margin, other two,
irregular in shape, in tornal and apical position;
cilia white; cilia line absent. Hindwing brownish;
cilia brownish.
Male genitalia (figs. 13-16). — Uncus deeply
indented, lobes slightly setose. Gnathos, lobes elon-
gate. Tegumen basally widening. Valva, sacculus
with long and distinct distal spine, costa strongly
convex in the middle, cucullus widely rounded.
79
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
7
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Figs. 9-12. Elachista megagnathos, holotype, male genitalia. 9-10, ventral view; 11, valva, lateral view; 12, aedoeagus,
lateral view. Scale 0.1 mm.
80
SRUOGA: New species of Elachistidae
Figs. 13-16. Biselachista spinigera, holotype, male genitalia. 13-14, ventral view; 15, lateral view: 16, juxta. Scale 0.1 mm.
81
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
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Figs. 17-24. Male genitalia. 17-19. Cosmiotes kopetdagica, holotype. 17, ventral view; 18, lateral view; 19, juxta, anellus
and digitate process. 20-24. Cosmiotes pallens, holotype. 20, ventral view; 21, valva, lateral view; 22, uncus, gnathos and
tegumen; 23, juxta, anellus and digitate process; 24, aedoeagus. Scale 0.1 mm.
82
Fig. 25. Distribution map: Perittia biloba (dot), Elachista
multipunctata (dot), Biselachista spinigera (triangle),
Cosmiotes kopetdagica (rectangle), Cosmiotes pallens
(rectangle).
Juxta, lobes broad, tapering dorso-lateral, small
group of setae on apical margin; digitate process
narrow at base, widest above the middle, setose.
Vinculum triangular. Aedeagus more or less
straight.
Biology. — Unknown. Holotype caught in Au-
gust.
Distribution (fig. 25). — Turkmeniya.
Cosmiotes kopetdagica sp. n. (figs. 17-19, 25)
Type material. — Holotype g: USSR, Turkmeniya,
western Kopet Dag, 40 km E Kara-Kala, 28.v.1988, V.
Sruoga (MG).
Diagnosis. — Closely related to C. amselt Parenti.
Distinguished from this species by the shape of the
juxta lobes and the very large gnathos.
External characters. — Male (female unknown).
Forewing length 3.6 mm. Frons and neck tufts
greyish white, whitish lustre. Antenna whitish grey
and brown ringed, distally slightly serrate. Labial
palp white with some lustre from above, brownish
from below. Thorax and tegulae brownish. Fore-
wing, ground colour brownish grey, some silvery
lustre, strongly mottled with brown scales, distal
part mottled by dark brown scales, distinct white
spots at tornus and costa, indistinct small whitish
spots near costa and approximately in the middle of
the forewing; cilia greyish; cilia line indistinct.
Hindwing greyish brown; cilia grey.
Male genitalia (figs. 17-19). — Gnathos very
large, oval, broadened. Valva, distal spine of saccu-
lus large, distinct, acute. Juxta, lobes short, triangu-
lar in ventral view, small setae; digitate process
long, distal part broad and setose. Vinculum trian-
gular; saccus long, gradually tapering. Aedeagus
long and slender, curved at base, long spine-like
cornuti.
Biology. — Unknown. Holotype caught in May.
Distribution (fig. 25). — Turkmeniya.
SRUOGA: New species of Elachistidae
Cosmiotes pallens sp. n. (figs. 20-24, 25)
Type material. — Holotype 4: USSR, Turkmeniya,
western Kopet Dag, 40 km E Kara-Kala, 28.v.1988, V.
Sruoga (MG).
Diagnosis. — Most similar to C. freyerella
(Hiibner). Easily distinguished from this and other
related species by more or less pale forewings,
broad gnathos and broad distal spine of sacculus,
large anellus and presence of cornuti in the aedea-
gus.
External characters. — Male (female unknown).
Forewing length 2.5 mm. Frons and neck tuft whit-
ish creamy. Antenna creamy brownish, distal part
finely serrate, scape brown, pecten creamy. Labial
palp creamy white. Thorax whitish to creamy
white, some yellowish-brown scales; tegulae brown.
Forewing, ground colour whitish to creamy white,
some yellowish-brown scales, brown scales in two
irregular spots in middle and abundantly present in
apical part of forewing, basal part of forewing paler,
dominated by whitish-creamy scales; cilia greyish-
creamy, some scattered brown scales; cilia line in-
distinct. Hindwing of holotype lost.
Male genitalia (figs. 20-24). — Gnathos broad.
Valva, distal spine of sacculus very distinct and
broad, not acute. Juxta, lobes short with fine setae,
apical margin sharp; digitate process long, distally
broad, setose; anellus very large. Vinculum with
long, narrow saccus. Aedeagus long, slender, with
acute cornuti.
Biology. — Unknown. Holotype caught in May.
Distribution (fig. 25). — Turkmeniya.
Fig. 26. Distribution map: Elachista fuscofrontella (trian-
gle), Elachista megagnathos (dot).
ACKNOWLEDGEMENTS
I wish to thank very much Dr. N. V. Savenkov
(Riga) for valuable information on Elachistidae of
Latvia. I am indebted to Dr. Rimantas Puplesis
(Minology working group, Vilnius) for his advice
85
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
and help as well as for making available the mater-
ial collected by him. Also I wish to express my
gratitude to R. Noreika (Minology working group)
and particularly to S. Jolanta, for their continuous
and kind help with my work.
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Received: 5 April 1989.
Revised version accepted: 1 February 1990.
BR. THEOWALD & P. OOSTERBROEK
Instituut voor Taxonomische Zoölogie UvA, Amsterdam
ZUR ZOOGEOGRAPHIE DER
WESTPALAARKTISCHEN TIPULIDEN.
IX. DIE TIPULIDEN DES VORDEREN ORIENTS.
Theowald, Br. & P. Oosterbroek, 1990. Zur Zoogeographie der Westpaläarktischen Tipu-
liden. IX. Die Tipuliden des Vorderen Orients. — Tijdschrift voor Entomologie 133: 85-
95, figs. 1-3, tabs. 1-4 + 1 appendix [ISSN 0040-7496] Published 31 July 1990.
A review is presented of the 198 species and subspecies of Tipulidae known from the Near
East. The Tipulidae fauna of the Near East distinctly belongs to the Westpalaearctic but has
an isolated position as is clear from the high amount of endemic species (67%).
The Near East can be divided in two regions of different faunal composition, the Southeast
with predominantly Mediterranean, and the Northwest with a majority of European and
Eurasiatic (sub)species.
About 50% of the (sub)species belongs to the subgenus Lunatipula. It is concluded that the
differentiation within the larger species-groups of Lunatipula dates back to at least the
Lower Miocene.
Both authors: Instituut voor Taxonomische Zoölogie (Zoölogisch Museum), Universiteit
van Amsterdam, Plantage Middenlaan 64, 1018 DH Amsterdam, Nederland.
Key words. — Diptera; Tipulidae; Near East; Zoogeography.
EINLEITUNG
In dieser Arbeit sind die Tipuliden des Vorderen
Orients zusammengefafst. Zum Vorderen Orient
sind hier gezählt: die asiatische Türkei (Anatolien),
die Sovjet Republiken südlich des Kaukasus (Trans-
kaukasien: Georgien, Azerbaidjan, Armenien),
Iran, Irak, Syrien, Libanon, Jordanien, Israel und
Nordägypten (Fig. 1). Untersuchungen der Tipuli-
denfauna dieser Gebiete haben erst spät angefan-
gen. Bis 1960 waren insgesamt etwa 50 Arten und
Unterarten bekannt, heute sind es 198. Damit ist
aber die Tipulidenfauna des Vorderen Orients si-
cher nicht vollständig bekannt. Manche neue Aus-
beute enthält wieder von dort unbekannte Arten
und von schon bekannten Arten neue Fundorte.
Für diese Arbeit wurden Daten aus der Literatur
und aus der Sammlung des Zoologischen Museums,
Amsterdam, zusammengetragen. Der Grofteil der
dort befindlichen Tipuliden des Vorderen Orients
wurde gesammelt von A. & U. Aspöck, H. v. d.
Brink, J. Lucas, H., T. & B. v. Oorschot, H. & R.
Rausch, F. Ressl und H. Wiering. Wir sind ihnen
und allen anderen Sammlern für das Material, das
sie uns überließen, sehr dankbar.
Besonders verplichtet sind wir unserem Freund
Günther Theischinger (Engadine, Australien) für
Ergänzungen und für das Überprüfen des deut-
schen Textes.
Für "Arten und Unterarten” ist im Text der
Kürze wegen die Bezeichnung “Arten” verwendet.
VEGETATION DES VORDEREN ORIENTS
Ursprünglich war der Vordere Orient (Rikli
1946; Atlas of the Arab World and the Middle East
1960) größtenteils trockene Grassteppe und Wüste,
Biotope, die kaum für Tipuliden geeignet sind. Es
gab aber auch Wald und Wiesen: hauptsächlich ent-
lang der Küsten, in den Gebirgen von Transkauka-
sien, Iran und Ostanatolien und entlang der
größeren Flüsse (z.B. Nil, Euphrat, Tigris) (Fig. 2).
In den höheren Lagen der Gebirge gab es große
Flächen von sommergrünem Laubwald und Misch-
wald, in den niederen Lagen mehr immergrünen
mediterranen Wald.
Der Vordere Orient ist eines der ältesten Kultur-
gebiete der Erde. Vor etwa 6000 Jahren wurde dort
bereits mit Landwirtschaft angefangen, und diese
hat sich seither über den größten Teil des Gebietes
ausgebreitet. Vor 3500 Jahren gab es schon viele
Städte. Eine so lange Zeit von Landwirtschaft und
andere Beeinflussung der Vegetation durch
menschliche Aktivitäten, wie zum Beispiel Holz-
nutzung für Bau und Brennstoff, hatte einen star-
ken Einfluß auf Flora und Fauna. Zuerst wurden die
feuchten Gebiete entlang der Flüsse für Landwirt-
85
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Balkan -
halbinsel È
Levantinisches
Becken
Schwarzes Meer
SudruBland
\ Kaspisches À Turk-
% Rers. Golf À
Meer \ menien
Fig. 1. Der Vordere Orient mit Hauptgebieten, Staten, tiirkischen Provinzen und Inseln. Von tiirkischen Provinzen
markiert # sind keine Tipuliden bekannt.
Hauptgebiet 1: Westanatolien
Le Lesbos 7 Bursa
4 # Istanbul 8 Kocaeli
5 Canakkale 9 Sakarya
6 # Balikesir 10 Bilecik
Hauptgebiet 2: Levantinisches Ktistengebiet
Kh Khios Cy Cyprus
Ik Ikaria 14 Izmir
Sa Samos 15 Aydin
Ko Kos 16 Mugla
Ro Rodhos 19 Antalya
Hauptgebiet 3: Zentralanatolien
22 # _ Eskisehir 28a Konya p. p.
27 Ankara 34 Kayseri
Hauptgebiet 4: Ostanatolien
32 # Gaziantep 50 Mardin
46 Sivas 51# Diyarbakir
47 # Malatya 52# Elazig
48 Adiyaman 53# Tunceli
49 Urfa 54 # _ Erzincan
Hauptgebiet 5: Nordanatolien
23 Bolu 39 # Corum
24 Zonguldak 40 Amasya
25 Kastamonu 41 Sinop
26 # Cankiri 42 # Samsun
Hauptgebiet 6: Georgien
Hauptgebiet 7: Azerbaidjan
Hauptgebiet 8: Armenien
Hauptgebiet 9: Iran
86
Kütahya
Usak
Manisa
Denizli
Icel
Hatay
Adana
Maras
Syrien
Nigde
Kirsehir
Erzurum
Bingöl
Kars
Agri
Mus
Tokat
Ordu
Giresun
Gümüsane
Burdur
Isparta
Afyon
Konya p. p.
Libanon
Israel
Agypten
Nevsehir
Yozgat
Bitlis
Van
Siirt
Hakkari
Trabzon
Rize
Artvin
schaft genutzt, und die ursprüngliche Vegetation
und Fauna ist dort fast restlos verschwunden. In den
feuchteren Gebieten der Gebirge wurden Laub- und
Mischwälder großtenteils abgeholzt, und der Boden
wurde über kürzere oder längere Zeit landwirt-
schaftlich genutzt. Im ganzen Gebiet sind heute nur
noch voneinander isolierte Laub- und Mischwald-
reste übrig, besonders in den weniger zugänglichen
Teilen der Gebirge. Viele der in älteren Zeiten für
Landwirtschaft genutzten Gebiete sind heute ent-
weder mit trockener mediterraner Hartlaubvegeta-
tion bewachsen, oder zu Steppen und Wüsten ge-
worden.
Die Vegetation des Vorderen Orients schließt im
Nordwesten und Norden nahe an jene der Balkan-
halbinsel und Südrußlands an. Von der Balkanhalb-
insel ist sie getrennt durch Bosporus und Dardanel-
len, die aber stellenweise weniger als 5 km breit
sind. Von Südrußland ist sie getrennt durch das
Schwarze Meer und den Kaukasus. Dieses Gebirge
ist bis über 5000 m hoch, fällt aber an den Küsten
des Schwarzen Meeres und des Kaspischen Meeres
bis ins Flachland ab. Im Nordosten ist der Vordere
Orient vom sibirischen Gebiet getrennt durch das
Kaspische Meer und die Steppen und Wüsten von
Turkmenien. Die Gebirge von Nordostiran gehen
aber allmählich in die von Afghanistan und Zen-
tralasien über. Für die mehr oder weniger feuchtig-
keitsliebenden Tipuliden ist somit im Norden des
Vorderen Orients Austausch mit der Balkanhalbin-
THEOWALD & OOSTERBROEK: Tipuliden
sel, mit Südrußland und mit Zentralasien wohl
schwierig, jedoch nicht auszuschließen. Im Süden
ist der Vordere Orient für die feuchtigkeitslieben-
den Tipuliden aber vollständig abgeschlossen vom
orientalischen und vom afrotropischen Gebiet
durch die Steppen und Wüsten von Südafghanistan,
Südiran, Irak, Syrien, Jordanien und Arabien. Fau-
nenaustausch mit diesen Regionen ist also nicht zu
erwarten.
ARTENLISTE (APPENDIX 1)
In der Artenliste sind die 198 Tipuliden des Vor-
deren Orients alphabetisch angeführt mit ihrer
Verbreitung innerhalb und außerhalb des Gebietes.
Aufgrund von Klima und Vegetation wurde der
Vordere Orient in 9 Gebiete aufgeteilt, wobei aus
praktischen Gründen soweit möglich die Grenzen
von Staaten und Provinzen benutzt worden sind
(Fig. 1). Die 9 Hauptgebiete sind: 1 Westanatolien,
2 das levantinische Küstengebiet, 3 Zentralanato-
lien, 4 Ostanatolien, 5 Nordanatolien, 6 Georgien,
7 Azerbaidjan, 8 Armenien, 9 Iran. Im Text zu Figur
1 sind die 9 Gebiete mit zugehörigen Staaten, tür-
kischen Provinzen und Inseln verzeichnet. Die
große türkische Provinz Konya ist aufgeteilt in den
feuchteren gebirgigen Westen und Süden (28) und
die mehr trockene Hochebene (28a). Die Inseln
wurden zum nächstgelegenen Festland gerechnet.
Nicht mitgezählt wurden Jordanien und Irak, von
Balkan - Schwarzes Meer
halbinsel
Levantinisches
Becken
SudruBland Turk -
menien
Kaspisches
Meer
=
Pers. Golf
Fig. 2. Die ursprüngliche Vegetation des Vorderen Orients, schwarz: feuchte Gebiete mit Wald und Wiesen, weiß:
trockene Gebiete mit Steppen und Wüsten.
87
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
welchen Staaten keine Tipuliden bekannt sind. Von
Syrien sind Tipuliden nur bekannt vom Küstenge-
biet, von Iran fast nur von den Gebirgen südlich des
Kaspischen Meeres. Von Ägypten ist nur eine Art
bekannt, der Kulturschädling T. (Tipula) orientalis.
In der Artenliste ist die Verbreitung über die 9
Hauptgebiete verzeichnet, und in Klammern, in
welcher der türkischen Provinzen oder auf welchen
Inseln. Ebenso sind die Verbreitungstypen im pa-
läarktischen Raum aufgenommen. Es bedeutet:
A Hauptverbreitung in Asien;
E Hauptverbreitung in Europa;
EA Hauptverbreitung in Eurasien;
M Nur verbreitet im mediterranen Raum der
Westpalaarktis.
Die Endemiten sind mit e bezeichnet und dahin-
ter steht jeweils einer der obengenannten Buchsta-
ben. Die großgeschriebenen Buchstaben weisen auf
die Verwandtschaft hin. So bedeutet zum Beispiel
eA: die Art ist endemisch im Vorderen Orient, und
ihre nächsten Verwandten haben asiatische Ver-
breitung.
ZOOGEOGRAPHISCHE BEMERKUNGEN
Stellung des Vorderen orients im
palaarktischen Raum
In Tabelle 1 sind die Tipuliden des Vorderen
Orients zusammengefaßt und geordnet nach ihrem
Vorkommen in den 9 Hauptgebieten und nach ih-
rer Verbreitung im paläarktischen Raum.
Der Vordere Orient beherbergt 8 Arten mit asia-
tischer Verbreitung oder Verwandtschaft (A und
eA), 45 mit europäischer (E und eE) und 17 mit
eurasischer (EA und eEA) Verbreitung oder Ver-
wandtschaft. Die eurasischen Arten gehören zur
westpalaarktischen Fauna, haben die Kaltzeiten am
Ende des Pleistozins im Balkan Refugium ver-
bracht und sich seither über Europa und über klei-
nere oder größere Teile Asiens ausgebreitet (Theo-
wald & Oosterbroek 1983, 1985).
Tabelle 1. Die Tipuliden des Vorderen Orients nach ih-
rem Vorkommen in den 9 Hauptgebieten.
Hauptgebieter MERS 475 CR SN OT
Verbreitungstyp
A O7 21 REDA 2 3
e tot or Oel OO) 3
E GM 3 7 PWG 9 DANS
eE NND PS2 1
EA 4 2 6 © 10, 7 5. 2
eEA 1 On it AL 2 3 5
M WUO 2e Foe Oor À SA
eM 16,59 maw. Se 19S SAS 107,
Total DI Bik WD 3b) Dik Gy) 410530578198
Insgesamt 128 Arten gehören zur mediterranen
Fauna der Westpaläarktis (M und eM). Mit 190 zur
westpaläarktischen Fauna gehörigen Arten, gegen-
über nur 8 ostpaläarktischen, gehört der Vordere
Orient deutlich zur Westpaläarktis, ganz im Gegen-
satz zum borealen Europa, wo 64% der Tipuliden
Arten von ostpaläarktischer Herkunft sind (Theo-
wald & Oosterbroek 1985).
Innerhalb der Westpaläarktis hat der Vordere
Orient jedoch eine isolierte Lage, was sich im hohen
Anteil der Endemiten (67%) und im hohen Anteil
der mediterranen Arten (64%) zeigt. In jedem ein-
zelnen der südlichen Gebiete Europas (Balkanhalb-
insel, italienisches Festland, iberisches Festland)
gibt es weniger als 40% Endemiten und weniger als
35% mediterrane Arten.
Faunenzusammensetzung der 9 Hauptgebiete
Es gibt große Unterschiede in der Zahl der Ar-
ten, die von jedem einzelnen der 9 Hauptgebiete
bekannt sind (Tabelle 1).
Zentralanatolien (Hauptgebiet 3) zählt nur 15
Arten. Dieses Gebiet ist eine trockene Hochebene
mit Steppen und Wüsten, zum Teil bewässert und
landwirtschaflich genutzt. Was Tipuliden betrifft,
ist es wahrscheinlich der ärmste Teil Anatoliens.
Von Iran (Hauptgebiet 9) kennen wir 28 Arten.
Von diesem Land sind aber relativ wenige Ausbeu-
ten bekannt, und wahrscheinlich kommen dort
mehr Arten vor und sind besonders noch ostpa-
läarktische Arten zu erwarten.
Es gibt keine deutlichen Hinweise, daß in einigen
der weiteren Gebiete auffallend mehr, in anderen
auffallend weniger gesammelt worden ist, obwohl
Unterschiede in der Zahl der bekannten Arten
deutlich sind. Wahrscheinlich hängen diese Unter-
schiede mit dem Aufbau der Landschaft zusammen.
Tabelle 2 gibt die prozentuelle Faunenzusam-
mensetzung der Hauptgebiete. In dieser Tabelle
sind die europäischen und eurasischen Tipuliden
zusammengezählt. Aufgrund der Faunenzusam-
mensetzung läßt sich der Vordere Orient in ein
südwestliches Gebiet (Hauptgebiete 1 und 2) und
ein nordöstliches (Hauptgebiete 4 bis 9) aufteilen.
Im südwestlichen Gebiet gibt es vorwiegend medi-
terrane Arten. Obwohl die europäischen und eura-
sischen Arten nur 31% der Gesamtfauna darstel-
len, sind sie im Nordosten in der Mehrheit.
Prozentuell sind die Unterschiede mehr als deut-
lich: im Südwesten 69-78% mediterrane und weni-
ger als 30% europäische Arten, im Nordosten 48-
67% europäische und weniger als 27-43% mediter-
rane Arten. Hauptgebiet 3, mit nur 15 Arten
schließt sich mit 60% europäischer Arten dem
Nordosten an.
In Tabelle 3 sind die Tipuliden des Vorderen
Orients eingeteilt in Arten mit südwestlicher Ver-
breitung, Arten mit nordöstlicher Verbreitung und
Arten, die im Südwesten und im Nordosten vor-
Tabelle 2. Prozentuelle Faunenzusammensetzung der
Tipuliden des Vorderen Orients in den 9 Hauptgebieten.
Hauptgebiete 1 2 3 4 5 6 7 8 9 Total
asiatische
Arten De DO Ged Be WS I Tevet
europäische
und eurasische
Arten 29 20 60 S1 58 58 48 67 54 31
mediterrane
Arten 69 78 40 43 39 38 40 27 39 65
kommen. Zu Arten mit südwestlicher Verbreitung
werden jene gezählt, die wenigstens in Westanato-
lien und/oder dem levantinischen Kiistengebiet
vorkommen. Einige von diesen reichen mehr oder
weniger weit in den Westen von Nordanatolien
oder in den Südwesten von Ostanatolien, sie fehlen
aber in Transkaukasien und Iran. Zu Arten mit
nordöstlicher Verbreitung sind jene gezählt, die in
Transkaukasien und Iran Vorkommen. Zum Teil
kommen sie auch im Osten von Nordanatolien und
in größeren Teilen Ostanatoliens vor. Sie fehlen
aber in Westanatolien und im levantinischen
Kiistengebiet.
Es gibt im Vorderen Orient von 198 Arten nur
19, die weitverbreitet sind. Sie sind ziemlich
gleichmäßig über das ganze Gebiet verbreitet. Es
fällt aber auf, daß 11 der 15 Arten von Zentralana-
tolien zu dieser Gruppe gehören. In den anderen
Gebieten sind sie deutlich in der Minderheit. Unter
ihnen sind mehrere bedeutende Kulturschädlinge.
Die übrigen 179 Arten sind ziemlich gleichmäßig
auf Südwesten (93) und Nordosten (86) verteilt.
Zwischen beiden Gebieten gibt es also nicht nur
einen Unterschied im prozentuellen Anteil von eu-
ropäischen und mediterranen Arten (Tabelle 2),
jedes der beiden Gebiete zählt überdies mehr als
80% charakteristische Arten, die im anderen Ge-
biet nicht vorkommen. Die beiden Gebiete haben
weniger als 20% ihrer Arten gemeinsam. Der Vor-
dere Orient läßt sich somit in zwei deutlich vonein-
ander verschiedene Faunengebiete einteilen: Süd-
westen und Nordosten. Ursache dafür ist wohl die
Tatsache, daß beide Gebiete schon längere Zeit
durch die Steppen und Wüsten von Zentralanato-
lien voneinander isoliert sind. Austausch von Arten
war nur möglich entlang der Küste des Schwarzen
Meeres über Nordanatolien und vielleicht auch
noch zwischen dem levantinischen Küstengebiet
und Ostanatolien im Süden der asiatischen Türkei.
Die europäischen und eurasischen Arten im Vor-
deren Orient
Die Mehrzahl der Arten mit europäischer oder
eurasischer Verbreitung (E, EA) haben die Kaltzei-
ten am Ende des Pleistozäns in einer Reihe von
Kleinrefugien auf der Balkanhalbinsel verbracht
THEOWALD & OOSTERBROEK: Tipuliden
und sich seither über Europa oder Eurasien ausge-
breitet (Theowald & Oosterbroek 1983, 1985). Von
den insgesamt 40 nichtendemischen europäischen
Arten im Vorderen Orient zeigen 31 eine ziemlich
geschlossene Verbreitung von der Balkanhalbinsel
über die Ukraine und Südrußland bis nach Georgien
und/oder Azerbaidjan, und von dort reichen sie
zum Teil weiter bis nach Iran oder in den Osten von
Nord- und Ostanatolien, 7 reichen sogar bis nach
Westanatolien und 4 bis in das levantinische Küs-
tengebiet. Nur 4 der 40 nichtendemischen europäi-
schen Arten sind von den Gebieten nördlich des
Schwarzen Meeres und vom östlichen Faunenge-
biet des Vorderen Orients unbekannt. Sie sind nur
bekannt vom Westen des Vorderen Orients. Die
Verbreitung von 5 Arten is unklar.
Ähnliches finden wir unter den 22 Endemiten
mit europäischer oder eurasischer Verwandtschaft
(eE, eEA). Die meisten haben eine nächstver-
wandte Art in den Gebieten nördlich des Schwarzen
Meeres und auf der Balkanhalbinsel. Insgesamt 16
dieser Endemiten sind im östlichen Faunengebiet
verbreitet, 2 davon bis Westanatolien. Rein westli-
che Verbreitung haben nur 6 Arten.
Die meisten europäischen und eurasischen Arten
(47 aus 62) zeigen also mehr oder weniger deutlich
eine Verbreitung von der Balkanhalbinsel über die
Gebiete nördlich des Schwarzen Meeres bis in das
östliche Faunengebiet des Vorderen Orients. Nur
15 Arten sind entweder nur vom westlichen Fau-
nengebiet bekannt oder Verbreitung und/oder Ver-
wandtschaft sind unklar. Die Gebiete nördlich des
Schwarzen Meeres waren jedenfalls im Saalien
(Rif) trockene Kältesteppen, und wenig weiter im
Norden befand sich die Eisdecke des Ural. Der
Osten des Vorderen Orients war damals also deut-
lich von der Balkanhalbinsel isoliert. Es ist deshalb
anzunehmen, daß die europäischen Tipuliden sich
erst rezent, am Ende des Pleistozäns oder im Ho-
lozän, von der Balkanhalbinsel über die Ukraine
und Südrußland bis nach dem Osten des Vorderen
Orients ausgebreitet haben. Ausbreitung vom Sü-
den der Balkanhalbinsel über Bosporus und Darda-
nellen bis nach Westanatolien hat anscheinend be-
deutend weniger stattgefunden.
Die mediterranen Arten im Vorderen Orient
Von den mediterranen Arten des Vorderen
Tabelle 3. Verteilung der südwest-, nordost- und weit-
verbreiteten Arten über die 9 Hauptgebiete.
kauptgebieten | DIESE CRA SOS Total
Verbreitung
südwest BO) Fl AO BO Or 0093
nordost OO OMS ECS 2 285162202286
I CRELO UL TLS NISSAN
815157357 517765740, 302287198
weitverbreitet
Total
89
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Tabelle 4. Artengruppen von Lunatipula nach den in Figur 3 erwähnten Gruppen und der Zahl der endemischen Arten
und Unterarten im mediterranen Gebiete (West Mediterr. inkl. Makronesien und Nordwest Afrika; Italien inkl. Korsika,
Sardinien und Sizilien; Balkan südlich Österreich und Ungarn; Vorderer Orient wie für diese Arbeit umschrieben). Die
Gezamtzahl der paläarktischen Lunatipula Arten und Unterarten ist 340.
Mediterrane Gebiete West Italien | Balkan Vordere Paläarktische
Mediterr. Orient Arten
Falcatae
falcata-Gruppe 9 3 14
helvola-Gruppe 1 3 5
bullata-Gruppe 4 5
10 10 — —
Fasciculatae A
livida-Gruppe 11 4 10 17 74
ill 4 10 17 (= 40%)
Fasciculatae B
verrucosa-Gruppe 5 6
caudispina-Gruppe 3 8 11
fascingulata-Gruppe 3 2 8 1 19
3 5 16 6 (= 20%)
Spinosae A
fascipennis-Gruppe 2 1 8
fulminis-Gruppe 1 3
truncata-Gruppe 18 22
— 2 20
Spinosae B1
caucasica-Gruppe 1 1 5 7
macroselene-Gruppe 1 1 4 19 30
peliostigma-Gruppe 2 2 12 19
1 4 7 35 (= 74%)
Spinosae B2
pustulata-Gruppe 5 7
clio-Gruppe 5 1 6
phaidra-Grupe 4 l 6
acuminata-Gruppe 3 13 33 53
lunata-Gruppe 1 8 14
9 — 22 43 (= 58%)
Orients sind 21 (M) unverändert auch von Gebieten
außerhalb des Vorderen Orients bekannt und sind
weiter 107 Endemiten (eM). Von den 21 Nichten-
demen sind 11 verbreitet vom Westen des Vorderen
Orients bis auf die Balkanhalbinsel (einige sogar bis
in das westmediterrane Gebiet), gegenüber nur 4,
die vom Osten des Vorderen Orients bis nach Siid-
rußland oder weiter verbreitet sind. Das bedeutet,
daß es mehr Austausch von mediterranen Arten
gegeben hat im Westen als im Osten. Eine Art (T.
(L.) cressa) kommt auf Rodhos und Kreta vor. Sie
gehört in die phaidra-Gruppe mit kretischer Ver-
breitung (siehe unten). Von 5 weiteren Arten ist die
Verbreitung unklar. Sie sind entweder über das
ganze ostmediterrane Gebiet, auch nördlich des
Schwarzen Meeres, verbreitet, oder sie zeigen in
ihrer Verbreitung große Disjunktionen.
Von den 107 Endemiten haben 60 eine südwes-
tliche, und 39 eine nordöstliche Verbreitung. Ob-
90
wohl die Arten also größenteils rein südwestlich
oder nordöstlich sind, gibt es unter den Arten-
gruppen, zu welchen sie gehören, kaum Hinweise
auf eine derartige geografische Zweiteilung. Die
größeren Artengruppen (/ivida-, macroselene-, pe-
liostigma-, acuminata- und lunata-Gruppe) haben
etwa gleichviele Arten im Westen und im Osten des
Vorderen Orients. Wahrscheinlich waren diese
Gruppen schon über den Vorderen Orient verbre-
itet, bevor dieses Gebiet durch Austrocknung des
zentralanatolischen Gebietes in zwei Faunenge-
biete geteilt wurde.
Die Untergattung Tipula (Lunatipula) im Vor-
deren Orient
Im Vorderen Orient gehören 124 der 198 Tipu-
liden (63%) zur Untergattung Lunatipula. Die pa-
läarktischen Arten dieser Untergattung wurden,
hauptsächlich durch Mannheims (1963-1968) und
Savtshenko (1964), eingeteilt in 27 Artengruppen,
von denen 19 auch im Vorderen Orient heimisch
sind. Die Verwandtschaftsbeziehungen zwischen
den Artengruppen wurden durch die beiden Auto-
ren aber nicht klargestellt.
Eine Einteilung der paläarktischen Lunatipula-
Arten in einige größere Hauptgruppen wurde
schon von Riedel (1913) versucht, der ohne weitere
Begründung die Untergattung aufteilte in Falcatae,
Fasciculatae und Spinosae. Es ist möglich diese Ein-
teilung teilweise durch Synapomorphien zu bele-
gen, miteine weiteren Aufspaltung der Fasciculatae
und Spinosae (Fig. 3).
l. Adminiculum mit deutlichen membranösen
Strukturen seitlich des Penisführers, die ihn größ-
tenteils oder ganz verdecken. Plesiomorph sind
derartige Strukturen nicht anwesend und liegt der
Penisführer unbedeckt.
2. Seitlich des Penisführers finden sich nebst die
differenzierte größere oder kleinere blattförmige
Strukturen überdies kleinere oder größere Dornen.
3. Seitlich des Penisführers finden sich stark
sklerotisierte Dornen und nur Reste von membra-
nösen blattförmigen Strukturen.
4. Am Hinterrand des 8. Sternits der Männchen
sind zwei Anhänge die meist kurze Dornen tragen.
5. Die Anhänge des 8. Sternits tragen einen lan-
gen geschwungenen Dorn.
6. Zwischen den Anhängen des 8. Sternits befin-
det sich ein Haarplättchen.
FASCICULATAE A
FASCICULATAE B
SPINOSAE BI
SPINOSAE B2
FALCATAE
SPINOSAE A
Fig. 3. Kladogramm der palaarktischen Hauptgruppen
von Lunatipula.
THEOWALD & OOSTERBROEK: Tipuliden
Tabelle 4 gibt die Einteilung der 19 Artengrup-
pen des Vorderen Orients unter den 6 Hauptgrup-
pen von Figur 3, und die Zahl ihrer mediterranen
Endemiten. Drei Arten der Balkanhalbinsel, die
Mannheims in die falcata-Gruppe stellt, werden
hier, conform mit Savtshenko, zu der fascingulata-
Gruppe gezählt, wohin sie aufgrund des Baues des
Adminiculum gehören.
Aus Tabelle 4 geht hervor, daß den hochdifferen-
zierten Spinosae Bl und B2 im Vorderen Orient die
größte Bedeutung zukommt. Von den zugehörigen
121 mediterranen Endemiten dieser Gruppen ha-
ben 78 (64%) dort ihre Verbreitung. Es ist anzu-
nehmen, daß die Spinosae Bl und B2 sich im Vor-
deren Orient differenziert haben, und daß sich
später Arten von dort weiter ausgebreitet haben.
Von den übrigen Hauptgruppen haben Fascicu-
latae B und Spinosae A vorwiegend balkanische
Verbreitung, sind die Fasciculatae A ziemlich
gleichmäßig über die ganze Paläarktis verbreitet
(besonders die /zvida-Gruppe) und findet sich die
älteste Gruppe der Falcatae nur im westmediterra-
nen Gebiet.
Die Differenzierung der Hauptgruppen im ost-
mediterranen Gebiet hat wahrscheinlich schon im
Unteren Miozän oder früher angefangen. Eine so-
genannte kretische Verbreitung über die südaegäi-
schen Inseln zeigt sich in der bemacula-Subgruppe
(livida-Gruppe, Fasciculatae A), der verrucosa-
Gruppe (Fasciculatae B), der cretis-Subgruppe
(macroselene-Gruppe, Spinosae Bl) und der
phaidra-Gruppe (Spinosae B2) (Theowald & Oos-
terbroek 1990). Eine derartige Verbreitung geht
wahrscheinlich zurück auf eine Zeit, in der es von
Südgriechenland über Kreta und Rhodos bis nach
Westanatolien eine Landbrücke gab. Diese Land-
brücke ist in der zweiten Hälfte des Miozäns zu
Inseln auseinander gefallen (Parent 1988). Dies
bedeutet, daß die meisten Hauptgruppen von
Lunatipula, auch die hochdifferenzierten Spinosae
B1 und B2, schon damals existierten.
ZUSAMMENFASSUNG
Unter den insgesamt 198 Tipuliden Arten im
Vorderen Orient sind 8 ostpaläarktischen gegen-
über 190 westpaläarktischen. Der Vordere Orient
gehört somit klar zur Westpaläarktis. Charakteris-
tisch für das Gebiet ist die große Zahl von Endemi-
ten und mediterranen Arten.
Der Vordere Orient läßt sich in zwei Faunenge-
biete aufteilen: ein südwestliches und ein nordöst-
liches. Beide haben nur wenige Arten gemeinsam.
Sie sind voneinander getrennt durch die zentralana-
tolischen Steppen und Wüsten.
Fast ein Viertel der Arten des Vorderen Orients
ist auch über die mehr nördlich gelegenen und küh-
leren Gebiete Europas verbreitet. Sie finden sich
vorwiegend im östlichen Faunengebiet des Vorde-
Oi
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
ren Orients und es ist anzunehmen, daß die meisten
europäischen und eurasischen Arten sich im Spät-
pleistozän oder im Holozän von der Balkanhalbin-
sel über Südrußland dorthin ausgebreitet haben.
Fast die Hälfte der Arten des Vorderen Orients
gehören zu hochdifferenzierten Artengruppen von
Lunatipula, einer Untergattung von Tipula, die sich
dort seit etwa dem Unteren Miozän entwickelt hat.
LITERATURVERZEICHNIS
Mannheims, B., 1963. 15. Tipulidae. — In: E. Lindner
(Ed.): Die Fliegen der palaearktischen Region, 3 (5) 1,
Lief. 238: 137-176.
Mannheims, B., 1965. Idem, Lief. 256: 177-212.
Mannheims, B., 1966. Idem, Lief. 267: 213-256.
Mannheims, B., 1967. idem, Lief. 270: 257-288.
Mannheims, B., 1968. Idem, Lief. 275: 289-320.
Parent, G. H., 1988. Esquisse biogéographique de la Crete.
— Linneana Belgica 9: 344-387.
Riedel, M. P., 1913. Die paläarktischen Arten der Dipte-
ren-Gattung Tipula L. — Abhandlungen des Vereins
für naturwissenschaftliche Erforschung des Nieder-
rheins, Krefeld 1: 1-122.
Rikli, M., 1943. Das Pflanzenkleid der Mittelmeerländer.
Erster Band. — H. Huber, Bern: 1-436.
Rikli, M., 1946. Idem. Zweiter Band: 437-1094.
Rikli, M., 1948. Idem. Dritter Band: 1095-1418.
Savtshenko, E. N., 1964. Tipulidae. — Fauna SSSR (New
Series), Diptera 2 (4) 89: 1-503.
Theowald, Br. & P. Oosterbroek, 1980. Zur Zoogeogra-
phie der westpaläarktischen Tipuliden. I. Die Tipuli-
den von Nordafrika. — Beaufortia 30: 179-192.
Theowald, Br. & P. Oosterbroek, 1981. Idem. II. Die Ti-
puliden der iberischen Halbinsel. — Beaufortia 31: 1-
20.
Theowald, Br. & P. Oosterbroek, 1982. idem. IV. The
Tipulidae of Corsica and Sardinia with a note on Do-
lichopeza fuscipes Bergroth. — Mitteilungen der
schweizerischen entomologischen Gesellschaft 55:
317-332.
Theowald, Br., C. Dufour & P. Oosterbroek, 1983. Idem.
III. Die Tipuliden der europäischen Tiefebenen. —
Bonner zoologische Beitrige 34: 371-394.
Theowald, Br. & P. Oosterbroek, 1984. Idem. V. Die ita-
lienischen Tipuliden. — Fragmenta entomologica
Roma 17: 245-291.
Theowald, Br. & P. Oosterbroek. 1985. Idem. VI. Die
Tipuliden der montanen, alpinen und borealen Ge-
biete. — Bonner zoologische Beiträge 36: 185-220.
Theowald, Br. & P. Oosterbroek, 1986. Idem. VII. Die
Tipuliden der Balkanhalbinsel. — Tijdschrift voor
Entomologie 129: 1-13.
Theowald, Br. & P. Oosterbroek, 1987. Idem. VIII. The
Tipulidae of Israel and adjacent regions. — Israel Jour-
nal of Entomology 20: 71-83.
Theowald, Br. & P. Oosterbroek, 1990. Idem. X. The
Tipulidae of the Eastmediterranean islands. — Bijdra-
gen tot de Dierkunde (in press).
IR
APPENDIX 1: Artenliste
Ctenophora
1. elegans Meigen, 1818: 2 (Sa)-E
2. guttata Meigen, 1818: 1 (28), 4 (67), 6-EA
3. magnifica Loew, 1869: 6, 9-eEA
4. ornata Meigen, 1818: 2 (29)-E
Dictenidia
5. bimaculata (Linnaeus, 1761): 5 (58), 6, 9-EA
Dolichopeza
6. fuscipes Bergroth, 1889: 2 (Sa)-M
7. graeca Mannheims, 1954: 4 (59), 5 (58)-M
8. hirsuticauda Savtshenko, 1968: 6, 7-eM
9. schabriari Theowald, 1978: 9-eM
Nephrotoma
cornicina-Gruppe
10. aculeata (Loew, 1871): 4 (64), 5 (40, 56), 6,
7-EA
11. appendiculata appendiculata (Pierre, 1919): 1
(21, 28), 3 (27), 4 (63, 64), 5 (25, 57), 6,7,8,
9-E
12. beckeri (Mannheims, 1951): 2 (Cy, 29, 30, Li,
Is), 4 (50)-eM
13. cornicina (Linnaeus, 1758): 1 (20, 21, 28), 2
(Sa, Cy, Li, 15), G45 35) MOD SIMCA)
51(25 5418) CAS NOLEA
14. eugeniae (Savtshenko, 1957): 6-eEA
15. gwestfalica (Westhoff, 1880): 2 (14)-E
16. minuscula (Mannheims, 1951): 2 (Cy, 30, Sy,
Li, Is)-eE
17. nasuta Oosterbroek, 1975: 1 (7, 28), 4
(64)-eM
18. quadrifaria quadrifaria (Meigen, 1804): 7-E
19. quadrifaria farsidica (Savtshenko, 1957): 9-eE
20. schaueffelei (Mannheims, 1964): 7, 9-eM
21. tenuipes (Riedel, 1910): 6-EA
22. theowaldi Oosterbroek, 1978: 1 (28), 2 (Ko,
Ro, 14, 15, 19, 29)-eE
crocata-Gruppe
23. analis (Schummel, 1833): 1 (21, 28), 3 (45,
37), 4 (59, 62, 63, 64), 5 (57), 6, 7, 8, 9-EA
24. croceiventris lindneri (Mannheims, 1951): 2
(Ko, 33), 3 (27; 34, 35), 4 (46, 61, 62, 63, 64,
Op), Of), D (C2, D1), FSE
25. nox (Riedel, 1910): 1 (28), 3 (35), 4 (48, 50,
59, 60, 61, 64, 65, 66, 67), 7, 8, 9-eEA
26. pratensis (Linnaeus, 1758): 5 (56), 6, 8-E
27. scalaris scalaris (Meigen, 1818): 1 (7, 21), 2
(Ik, Ro, Cy, 14, 19, 30, Sy, Li, Is), 3 (34), 4 (50,
594035.64,63), SACS SC) OA SS I-ER
28 scalaris terminalis (Wiedemann, 1830):
5 (58), 6, 7, 8, 9-A
Nigrotipula
29. nigra (Linnaeus, 1758): 7-EA
Tipula
T. (Acutipula)
30. cypriensis Vermoolen, 1983: 2 (Cy)-eM
31. fulvipennis fulvipennis De Geer, 1776:
4 (59), 5 (58), 6, 7, 9-EA
32. fulvipennis nigroantennata
1961: 5 (45, 58), 6, 7, 9-eEA
33. isparta Vermoolen, 1983: 1 (20), 2
(Cy, 29)-eM
34. irrequieta Alexander, 1935: 7, 8,9-A
35. libanica Vermoolen, 1983: 2 (Li, Is)-eE
36. macra Savtshenko, 1961: 7, 9-eM
37. transcaucasica
1961: 8, 9-M
38. transcaucasica latifurca Vermolen, 1983: 1
CAS) 2) dksSankKomlOns le Te) A (C2; 164;
65, 67), 5 (56, 57)-M
Savtshenko,
transcaucasica Savtshenko,
T. (Beringotipula)
39. unca Wiedemann, 1817: 6, 7, 8-EA
T. (Dendrotipula)
40. flavolineata Meigen, 1804: 6, 7, 8-E
T. (Emodotipula)
41. saginata Bergroth, 1891: 4 (59), 8, 9-E
T. (Lunatipula)
acuminata-Gruppe
42. angela Mannheims, 1963: 2 (Li, Is)-eM
43. angelica Theowald, 1957: 2 (Sy, Li, Is)-eM
44. artemis artemis Theischinger, 1977: 2 (Kh,
Ik, Sa)-M
45. artemis asiaeminoris Theischinger, 1982: 2
(14, 15)-eM
46. auriculata Mannheims, 1963: 2 (31)-eM
47. berytia Mannheims, 1963: 2 (Li, Is)-eM
48. borysthenica Savtshenko, 1952: 1 (28)-M
49. brinki Theischinger, 1987: 2 (19)-eM
50. canakkalensis Theischinger, 1987: 1 (5)-eM
51. curvata Theischinger, 1977: 2 (31)-eM
52. cypris Mannheims, 1963: 2 (Ro, Cy)-eM
55. decolor Mannheims, 1963: 1 (7, 9)-M
54. dedecor Loew, 1873: 2 (Sa, Ko)-eM
55. emmahelene Theischinger, 1980: 2 (29)-eM
56. franzressli Theischinger, 1982: 2 (16)-eM
57. geja Savtshenko, 1968: 6-eM
58. horsti Theischinger, 1982: 1 (18), 2 (19)-eM
59. huberti Theischinger, 1982: 2 (19)-eM
60. hyrcana Savtshenko, 1973: 7-eM
61. istriana Erhan & Theowald, 1961: 1 (Le, 9, 10,
18, 21, 28)-M
62. kaplani Theowald & Oosterbroek, 1987:
2 (Is)-eM
63. kreissli Theischinger, 1987: 5 (25)-eM
64. leeuweni Theischinger, 1982: 2 (Sy, Is)-eM
65. macropyga Savtshenko, 1952: 8, 9-eM
66. montifer montifer Theischinger, 1977:
2 (31)-eM
THEOWALD & OOSTERBROEK: Tipuliden
67. montifer tasucuensis Theischinger, 1987:
2 (29)-eM
68. musensis Theischinger, 1987: 4 (63)-eM
69. osmana Mannheims, 1963: 1 (7), 5 (40), 6,
7-eM
70. parasimurg Savtshenko, 1968: 7-eM
71. peteri Theischinger, 1979: 1 (7)-eM
72. simurg Savtshenko, 1964: 7-eM
73. subacuminata Mannheims, 1963: 2 (29, 31,
33), 4 (60, 64)-eM
74. suleika Mannheims, 1963: 7, 9-eM
75. theowaldi Savtshenko, 1964: 6-eM
76. turca Mannheims, 1963: 1 (7)-eM
77. valerii Savtshenko, 1968: 6-eM
78. vermooleni Theischinger, 1987: 2 (Ko)-eM
adusta-Gruppe
79. adusta lucistriata Mannheims & Savtshenko,
1967: 2 (31), 4 (61)-A
caucasica-Gruppe
80. caucasica Riedel, 1920: 6-eM
81. kinzelbachi Theischinger, 1982: 2 (Sy)-eM
82. quadridentata paupera Savtshenko, 1964:
6-eM
83. talyshensis Savtshenko, 1964: 7-eM
clio-Gruppe
84. chelifera Savtshenko, 1964: 4 (59), 5 (58),
6-eM
fascingulata-Gruppe
85. eugeniana Simova-Tosic, 1972: 6-M
86. praecox Loew, 1873: 6-M
87. rbynchos Theischinger, 1977: 2 (14)-eM
fascipennis-Gruppe
88. fascipennis Meigen, 1818: 6-E
helvola-Gruppe
89. helvola Loew, 1873: 1 (Le, 7, 28), 2 (Sa, 16, 19,
Is), 5 (56)-E
laetabilis-Gruppe
90. humilis Staeger, 1840: 6-E
livida-Gruppe
91. adzharolivida Savtshenko, 1968: 6-eM
92. bimacula bimacula Theowald, 1980: 1
(Le), ?9-M
93. cinerella Pierre, 1919: 1 (20, 28), 2 (19)-M
94. deserticola Savtshenko, 1968: 7-eM
95. freidbergi Theowald & Oosterbroek, 1987:
2 (Is)-eM
96. kumerloevi Mannheims, 1968: 2 (31)-eM
97. kybele kybele Mannheims, 1968: 1 (28), 2 (29,
Li, Is), 3 (35), 4 (63, 64, 66, 67)-eM
98. kybele russula Theischinger, 1977: 1 (18, 20,
28), 2 (19)-eM
93
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
99. livida morosa Savtshenko, 1964: 6-eE
100. maija Savtshenko, 1973: 6-eM
101. mendli Martinovsky, 1976: 1 (Le, 7, 17, 21),
2 (Kh, Ko, Sa, 16)-eM
102. nigdeensis Bischof, 1905: 3 (28a, 35)-eM
103. pseudowolfi Theischinger, 1979: 1 (28), 2 (Sa,
16)-eM
104. rbodolivida Theowald, 1972: 2 (Ko, Ro, 16,
19)-eM
105. saltatrix Savtshenko, 1964: 6-eM
106. sternalis Theischinger, 1977: 2 (Is), 4
(64)-eM
107. trapeza Theischinger, 1982: 1 (21)-eM
108. turcolivida Mannheims, 1968: 2 (31)-eM
109. vulpecula Theischinger, 1979: 4 (65), 5
(56)-eM
lunata-Gruppe
110. furcula Mannheims, 1964: 1 (5, 7, 21, 28), 2
(Kh, Ko, Ro, 14, 15, 16)-M
111. Aybrida hybrida Savtshenko, 1952: 7-eM
112. hybrida altivolans Savtshenko, 1971: 6-eM
113. lunata Linnaeus, 1758: 6, 7, 8-E
114. paravelox Theischinger, 1987: 2 (33)-eM
115. pseudolunata pseudolunata Theischinger,
1980: 1 (21, 28)-eM
116. pseudolunata spinalonga Theischinger, 1982:
6-eM
117. soost soost Mannheims, 1954: 1 (Le, 5, 7, 18,
2028) Sa, Ro 1415416419293),
3 (34, 35), 6-M
118. soosz izmirensis Theischinger, 1987: 2 (14)-
eM
119. subvelox Savtshenko, 1968: 5 (25), 6-eM
120. wmicornis Theischinger, 1977: 5 (25, 43)-eM
macroselene-Gruppe
121. acudens Theischinger, 1977: 2 (33)-eM
122. adapazariensis Theischinger, 1987: 1 (8)-eM
123. aurita Riedel, 1920: 6, 7, 8-eM
124. carens Theischinger, 1987: 1 (8)-eM
125. cedrophila Mannheims, 1963: 1 (10), 2 (Ro,
Li, Is)-eM
126. christophi Theischinger, 1982: 2 (Sa, 14)-eM
127. dumetorum Savtshenko, 1964: 6-eM
128. holzschuhi Theischinger, 1977: 9-eM
129. zliensis Mannheims, 1965: 2 (31)-eM
130. zmbecilla Loew, 1869: 2 (Ro)-eM
131. kerkis Theischinger, 1977: 1 (21)-eM
132. lithophila Savtshenko, 1968: 7-eM
133. /ucasi Theischinger, 1987: 4 (67)-eM
134. oorschotorum Theischinger, 1987: 1 (21)-eM
135. palifera Mannheims, 1965: 2 (19, 33), 3 (35)-
eM
136. rutila Savtshenko, 1952: 2 (33), 8-eM
137. selenis Loew, 1873: 2 (Ro)-eM
138. subbispina Savtshenko, 1952: 9-eM
139. tibonella Theischinger, 1977: 1 (Le, 11), 2
(14, 19)-M
94
140. zaitzevi Savtshenko, 1952: 6, 8-eM
mellea-Gruppe
141. affinis Schummel, 1833: 6-E
peliostigma-Gruppe
142. antilope Theischinger, 1977: 4 (67), 9-eM
143. astigma Savtshenko, 1968: 6-eM
144. didymotes Theischinger, 1977: 4 (61, 64, 65,
67)-eM
145. micropeliostigma Mannheims, 1965: 1 (21)-
eM
146. ornithogona Theischinger, 1982: 2 (16)-eM
147. peliostigma peliostigma Schummel, 1833: 1
(US) 2A (Sa xls ID) ZSNSTI NOMAD E
148. peliostigma burdurafyonensis Theischinger,
1987: 1 (18, 21)-eE
149. pseudopeltostigma Mannheims, 1965: 2 (Li,
Is)-eM
150. renate Theischinger, 1982: 1 (18)-eM
151. selene Meigen, 1830: 6-E
152. semipeliostigma Mannheims, 1965: 2
(31)-eM
153. strigosa Savtshenko, 1952: 5 (58), 6-eM
154. ulrike Theischinger, 1982: 1 (28)-eM
155. wrswlae Mannheims, 1965: 2 (19)-eM
phaidra-Gruppe
156. cressa Mannheims, 1965: 2 (Ro)-M
157. sciurus Theischinger, 1977: 1 (20, 28), 2 (19,
31)-eM
truncata-Gruppe
158. arnoldu Savtshenko, 1957: 6-M
verrucosa-Gruppe (= brunneinervis-Gruppe)
159. dracula Theischinger, 1977: 1 (21), 2 (Kh, Ko,
16)-eM
160. neutra Theischinger, 1982: 1 (28), 2 (19)-eM
161. quinquespinis Theischinger, 1980: 1 (Le, 17,
18, 28), 2 (16, 19, 29)-eM
162. teunisseni Theischinger, 1979: 2 (Ro)-eM
163. verrucosa verrucosa Pierre, 1919: 6, 7, 8-M
164. verrucosa sinedente Theischinger, 1980: 1
CUS A 23) AGO) 3) (AID), A (EO), SIE
43, 55)-eM
zimini-Gruppe
165. zimini semiopaca Savtshenko, 1964: 7-A
T. (Mediotipula)
166. anatoliensis Theowald, 1978: 1 (28)-eE
167. caucasiensis Theowald, 1978: 6-eE
168. stigmatella Schummel, 1833: 1 (7), 8-E
T. (Pterelachisus)
169. luteobasalis luteobasalis Savtshenko, 1964: 4
(64), 5 (58), 6, 8, 9-eE
170. luteobasalis alata Theowald, 1980: 4 (64)-eE
IA
trichopleura Savtshenko, 1964: 6-eE
T. (Savtshenkia)
172.
17,5
174.
75
176.
7/7.
178.
179:
180.
181.
baltistanica Alexander, 1935: 6, 7-A
interserta Riedel, 1913: 6, 7, 8-E
jeekeli Mannheims & Theowald, 1959: 1 (Le),
2 (Ro)-M
nivalis Savtshenko, 1961: 6-eEA
obsoleta Meigen, 1818: 5 (56), 6-E
ornata Theowald & Oosterbroek, 1987: 2
(Is)-eE
pechlaneri Mannheims & Theowald, 1959:
9-E
rufina rufina Meigen, 1818: 2 (Cy, Li, Is),
4 (67), 7, 9-E
serrulifera Alexander, 1942: 5 (58), 6-E
subalpium Savtshenko, 1961: 6-eE
T. (Schummelia)
182.
183.
ahrenst Savtshenko, 1957: 6-eE
varicornis Schummel, 1833: 8-E
T. (Tipula)
184.
185.
186.
italica errans Theowald, 1984: 1 (Le), 5 (56),
7, 8, 9-M
oleracea Linnaeus, 1758: 2 (Ro)-E
orientalis Lackschewitz, 1930: 1 (Le, 5, 18, 20,
DIN SNN NRO el 4 15129 Sv
Is, Ag), 3 (34), 4 (46, 48, 59, 63, 64, 65), 5 (41,
44, 58), 6, 7, 8, 9-M
187.
THEOWALD & OOSTERBROEK: Tipuliden
subcunctans Alexander, 1921: 8-EA
T. (Vestiplex)
188.
189.
190.
LOE
192:
kosswigi Mannheims, 1953: 1 (7)-eA
pallidicosta pullata Savtshenko, 1960: 1 (28),
4 (61), 5 (56), 6, 7, 8-eE
relicta Dia & Theowald, 1982: 2 (Li, Is)-eA
semivittata semivittata Savtshenko, 1960:
5 (45, 58), 6, 7-E
semivittata dissimilis Savtshenko, 1964: 6-eE
T. (Yamatotipula)
193.
194.
195.
196.
197.
198.
caesia Schummel, 1833: 1 (17, 21, 28), 3 (27),
4 (59, 63, 64), 6-E
caucasimontana Savtshenko, 1955: 4 (67),
7-eA
lateralis lateralis Meigen, 1818: 1 (7, 17, 18,
ZO W228) 82 (Sato Ko Ro 942993933)
Sy, Li, Is), 3 (27), 4 (46, 49, 50, 59, 60, 64, 67),
5 (24, 56), 6, 8-E
lateralis tranensis Theowald, 1978: 4 (64), 9-
eE
pruinosa Wiedemann, 1817: 6-EA
submontium Theowald & Oosterbroek, 1981:
INNEN IES TA ICE
8-E
Received: 23 March 1990
Revised version accepted: 16 May 1990
95
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
BOOK REVIEW
Johansson, R., E. S., Nielsen, E. J. van Nieukerken and B.
Gustafsson, 1989. The Nepticulidae and Opostegidae
(Lepidoptera) of North West Europe. - Fauna Ento-
mologica Scandinavica 23: 1-739, text-figs. 1-111, ta-
bles 1-3, figs. 1-1122. [ISBN 90 04 08698 6 (2 parts),
E. J. Brill, Leiden, The Netherlands. Price 220 Dutch
Guilders/US $ 110.-].
For the first time an up to date treatment is given
for all the 121 species of Nepticulidae and four
species of Opostegidae occurring in Fennoscandia,
Denmark, the British Isles, The Netherlands, Bel-
gium, France north of Paris, West and East Ger-
many, Poland and the Baltic states. This two vo-
lume work figures in full colour all species and gives
additional colour illustrations in cases of sexual
dimorphism or polymorphism. The colour illustra-
tions are of very high quality and make the book
even more a must for specialists, lepidopterists in
general and book-lovers alike. Line drawings of
male and female genitalia and leaf-mines for all
species are given. Also the larvae of most species
are figured.
Identification of the species is facilitated by var-
ious keys: key to the tribes and families of Nepti-
culoidea; key to the mines and larvae, arranged per
host plant genus; keys to the males, based on exter-
nal characters; keys to the males, based on genitalia;
keys to the females, based on external characters
and keys to the females, based on genitalia. Unfor-
tunately the key for the females of Nepticulini (the
first 76 species) based on the genitalia would be too
much a copy of the full descriptions and has been
omitted. But upon checking the line drawings of the
female genitalia a definite identification is easily
reached.
Each species is fully redescribed and biology and
distribution given. The chapters on immature
stages, biology, collecting and rearing, will undoubt-
edly stimulate many people to keep an eye out for
the mines of the Nepticulidae.
The phylogeny, classification and nomenclature
are briefly, but adequately, dealt with in a separate
chapter. One minor point to be mentioned in the
section which deals with the Nepticulini is the lack-
ing of the flight periods for the adults and the
months in which the larvae can be found. This in
contrary to the other part of the book where this
information is given. This book has a very high
scientific standard, but is easily accessible for the
non-specialist at the same time.
[R.T.A. Schouten]
96
J. vAN TOL
National Museum of Natural History, Leiden
KEY TO THE MALESIAN SPECIES OF LEPTOGOMPHUS
SELYS, WITH THE DESCRIPTION OF A NEW SPECIES
FROM SABAH (ODONATA, GOMPHIDAE).
Descriptions and records of Malesian Odonata, 1.
Tol, J. van, 1990. Key to the Malesian species of Leptogomphus Selys, with the description
of a new species from Sabah (Odonata, Gomphidae). Descriptions and records of Malesian
Odonata, 1. — Tijdschrift voor Entomologie 133: 97-105, figs. 1-37, tabs 1-2 [ISSN 0040-
7496]. Published 31 July 1990.
A new species of Leptogomphus, L. pasia, based on one female from S. Sabah (Northern
Borneo), is described. A congeneric and presumably conspecific male from E. Sabah is also
described. A key to the eleven species of this genus recorded from Malaysia, Indonesia, the
Philippine Islands, and the adjacent areas in Burma and Thailand, is included. Many
characters are illustrated for the first time.
J. van Tol, National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The
Netherlands.
Key words. — Leptogomphus; key; Malesia; new species.
INTRODUCTION
During the 1987 expedition of the Rijksmuseum
van Natuurlijke Historie (Leiden) to Sabah, north-
ern Borneo, a wealth of interesting Odonata was
collected. In this paper a new species of Lepto-
gomphus Selys is described, based on a single fe-
male. It was collected in the vicinity of Long Pasia,
a village still surrounded by vast areas of virgin
rainforest. This area, situated close to the borders
of Sarawak and Kalimantan at c. 1000 m above sea
level, is poorly investigated for the aquatic fauna up
to now. The Odonata fauna seems to be most closely
related to those of Mount Dulit and other higher
peaks in Sarawak. This may be exemplified by re-
cords of Coeliccia cyaneothorax Kimmins, formerly
only known from Mt. Dulit, and Burmagomphus
insularis Laidlaw, of which only the holotype from
Sarawak was known. A complete list of the species
encountered will be published in due time.
SYSTEMATIC PART
Leptogomphus pasia spec. nov. (Figs. 1-6, 8, 16,
32-33)
Type material. — Holotype female: 'RMNH
Leiden/N. Borneo, S. SABAH. Beaufort, 105 km S
of: Long Pasia area. Sungai Pasia near confluence
with Sungai Maga. Rockpools and stream. Lower
montane evergreen tropical rainforest. 1250 m.
4°26’N 115°40’E. 3 April 1987. Leg. C. van Ach-
terberg’, in Museum Leiden (RMNH).
Other material. — Sabah, 60 km W of Lahad
Datu: Danum Valley Field Centre, 4°58’N
117°48’E. 2 May 1987, 1 male (J. van Tol) in
RMNH.
Description
Female. — Closely related to Leptogomphus
lansbergei and L. semperi, but immediately recog-
nizable by its extremely long vulvar lamina (vulvar
scale) (authors of species-group names in Lepto-
gomphus are given in table 1).
Head. — Depressed; frons, clypeus and labrum
protruding. Labium with prementum dark brown
to brownish black, lateral lobes greenish yellow,
broadly bordered with brownish black, especially
distally; movable hooks brownish black, central
part of mentum pale brown, laterally bordered with
creamish white. Mandibles reddish brown-black,
but visible part beside the labrum mostly greenish
yellow. Labrum black with two relatively small sub-
triangular pale spots close to the clypeus. Clypeus
black, with a paired oval depression on postclypeus,
together approximately as wide as clypeus between
the emarginate anterior side. Frons distinctly
swollen with a broad greenish yellow transversal
stripe, covering nearly half of the surface, pale
stripe interrupted in the middle by a dark triangular
97
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 1
Species included in the key of the Leptogomphus species of Malesia, with original references
Species Author; date: pages
L. coomansi Laidlaw 1936: 267-269
L. gestroi Selys 1891: 476-477
L. inclitus Selys 1878: 444 (sep 39)
L. lansbergei Selys 1878: 446 (sep 41)
L. lansbergei assimilis Kriiger 1899: 308
L. mariae Lieftinck 1948: 249-251
L. palawanus Asahina 1968: 364-365
L. pendleburyi Laidlaw 1934: 555-556
L risi Laidlaw 1933: 95-96
L. semperi Selys 1878: 443 (sep 38)
L. williamsoni Laidlaw 1912: 94-95
Original genus
Leptogomphus
Leptogomphus
Gomphus (Leptogomphus)
Gomphus (Leptogomphus ?)
Leptogomphus
Leptogomphus
Leptogomphus
Leptogomphus
Leptogomphus
Gomphus (Leptogomphus)
Leptogomphus
Type depository
RMNH / H male
MCSN / H male
BMNH / L female
IRSN + SMTD / S female
ZMPA ? / S male + female
RMNH / H female
CASC / H male
BMNH / H male
BMNH / H male
IRSN / H male
BMNH / H male
Abbreviations: BMNH = British Museum (Natural History), London; CASC = California Academy of Sciences, San
Francisco; IRSN = Institut Royal des Sciences Naturelles de Belgique, Bruxelles; MCSN = Museo Civico di Storia
Naturale, Genoa; RMNH = National Museum of Natural History, Leiden; SMTD = Staatliches Museum für Tier-
kunde, Dresden; ZMPA = Zoological Museum Polish Academy of Sciences, Warszawa. - H = holotype, L = lecto-
type, S = syntype.
Note: The type of L. assimilis should be selected from material in the Dohrn collection, which was deposited in Stet-
tin (now Szczecin, Poland); some parts of the collection of this museum have been transferred to ZMPA.
emargination. Vertex shiny black, but dullish black
behind the tubercles posterior to the lateral ocelli;
occiput also dull black, tubercles behind lateral ocelli
consisting of a larger pair in the centre between the
ocelli and a smaller pair latero-posteriorly of the
former; occiput without tuberculi (fig. 2), which are
usually present in congeneric species (e.g. L.
semperi, L. coomansi, L. mariae and L. williamsont)
(see also Lieftinck 1948, plate 8); occiput poste-
riorly with a distinct triangular emargination.
Thorax (for nomenclature of thoracic markings,
cf. Williamson 1907: fig. 29 (1)). - Prothorax with
anterior lobe erect, black, but greenish yellow in
lateral edges; middle lobe with deep median depres-
sion anteriorly. Synthorax (fig. 5) brownish black
with pale markings yellow; dorsal thoracic stripe
connected with mesothoracic half collar (as in L.
coomansı), antehumeral stripe narrow, somewhat
wider at posterior side; pale markings on mesepi-
meron and metepisternum more or less as in L.
lansbergei, but pale area on metepimeron much
smaller than in L. lansbergei (the dark stripe under
the pale area approximately of equal width). Legs
stoutly built; hind femora with two rows of 9-12
distinct short spines on innerside; all tibiae with
two rows of 9-10 long spines on innerside. Wings:
forewing 37 mm, hind wing 35 mm. Forewing with
16 Ax and 13 Px; hindwing with 12 Ax and 13 Px.
Hind wing with one cubito-anal crossvein; an in-
complete crossvein proximal to first primary Ax
present; distal costal brace closer to proximal costal
brace than to nodus; triangle, supratriangle and
subtriangle all without crossveins. Pterostigma 3.5
mm.
Abdomen. — Length 39 mm. Dark brown, nearly
black, except for a yellow middorsal stripe, which is
rather wide on segments 1-2, but much narrower to
98
nearly indiscernable on segments 3-8; lateral side of
segment 1 dark yellow, lateral side of segment 2 and
antero-lateral side of segment 3 also dark yellow.
Auriculae hardly developed, yellow. Vulvar lamina
diagnostic (figs. 3-4), extremely elongate, the tip
reaching posterior margin of segment 10, very
slender, the top acutely v-shaped.
Male. - As female (figs. 6, 16), but pale spots on
labrum larger than in female, subquadrangular (fig.
8); hind femora with many short spines on inner-
side instead of the rows of medium-sized spines as
in the female; wings with anal triangle three-celled,
crossveins meeting in one point; secondary genita-
lia and anal appendages very similar to L. lansber-
gei, but superior appendage of /ansbergez in dorsal
view curved outwards more conspicuous than in
pasia (figs. 30, 32); base of superior appendage of
pasia bulgy on outer margin; consequently, laterov-
entral ridge with spines along the outer margin of
the superior appendage nearly straight in lansber-
get and distinctly curved in pasta (figs. 31, 33).
Although I consider this male conspecific with
the female described above, I prefer to include only
the female in the type series of this species.
Differential diagnosis. - The female of this spe-
cies, superficially looking as L. lansbergei, is most
easily distinguished from its congeners by the pe-
culiar shape of the vulvar lamina. Also the male is
very similar to L. lansbergei, but can be distin-
guished from most species by the thoracic mark-
ings, and from lansbergei by the shape of the super-
ior appendages.
Etymology. - Pasia from the type locality Long
Pasia. To be considered as a noun in apposition.
Discussion
Up to now the following species of the genus
Leptogomphus were known from Borneo: L. coo-
mansi, L. martae, L. pendleburyi and L. williamsoni.
There is still some doubt about the occurrence of L.
lansbergei assimilis on Borneo, of which a male was
recorded from Bettotan, 4 August 1927 by Laidlaw
(1931). This specimen is not in the collection of the
British Museum (Natural History) (Brooks in litt.).
Also L. semperi, known from Mindanao, should be
VAN TOL: Malesian species of Leptogomphus
taken into account when considering the fauna of
Borneo. Earlier records by Martin (1904) and Wil-
liamson (1907) were considered erroneous by Laid-
law (1936). This species was also attributed to the
lansbergei Formenkreis by Laidlaw (1936).
L. pasia shows some affinity to L. coomansi and
L. mariae, sharing the connection between the dor-
sal stripe and the mesothoracic half collar (cf. Lief-
tinck 1948, plate 8). The absence of spiny tubercles
on the occipital plate, and the peculiar shape of the
Figs. 1-7. Leptogomphus from northern Borneo. - 1, L. pasta female holotype, head in frontal view; 2, Idem, hind margin
of occiput in dorsal view; 3-4, Idem, last abdominal segments in ventral and left lateral view; 5, Idem, synthorax in left
lateral view; 6, L. cf. paria, male from Danum Valley, synthorax in left lateral view; 7, L. pendleburyi, male holotype,
synthorax in left lateral view.
99
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 8-14. Leptogomphus. - 8, L. cf. pasia, male from Danum Valley, head in frontal view; 9, L. pendleburyi, male
holotype, idem; 10, L. lansberget assimilis, male from S. Sumatra, idem; 11, L. inclitus, female lectotype, head in frontal
view; 12, L. lansbergei assimilis, female from S. Sumatra, last abdominal segments in ventral view; 13, L. inclitus, female
lectotype, idem; 14, L. inclitus, female lectotype, hind margin of occiput in dorsal view.
100
vulvar scale are diagnostic characters for the pres-
ently described species. As already mentioned
above, it is, however, most closely related to L.
lansbergei from Java and Sumatra.
KEY TO THE LEPTOGOMPHUS SPECIES OF MALESIA
In the next key I have summarized the characters
to distinguish the species of Leptogomphus occur-
ring in Malesia (i.e. the Malay peninsula, Indonesia,
the Philippine Islands and the island groups north-
east of New Guinea). I have added several species
recorded from Burma and Thailand that might oc-
cur in this area. All species mentioned in the key are
listed in table 1 with the author and a reference to
the original publication. Leptogomphus svibleri
Asahina, however, recorded from Burma and Thai-
land, is not included in the key. The absence of an
incomplete basal cross-vein (Asahina 1970: 116) is
an indication that this species is not a
Leptogomphus in the present sense. Asahina
(1986: 17) has placed it in Heliogomphus. The
genus Leptogomphus Selys as defined by Carle
(1986) forms with Africogomphus Fraser and
Heliogomphus Laidlaw the tribe Leptogomphini in
the Epigomphinae. Characters to distinguish
Leptogomphus and Heliogomphus include lamina
on fore tibia of male (present/absent, respectively),
basal subcostal crossvein (present/absent), cells of
anal triangle in male (meet in one point/serially),
lamina anterior (raised transverse-platelike/bilo-
bate), superior appendages of male (more or less
straight/lyrelike).
The key is mainly based on characters in the
markings of the specimens, predominantly of the
head and the thorax. Colour pattern is a reliable
character in this group and the interpretation of
Table 2
Species (
coomansi
gestroi
inclitus
lansbergei lansbergei
lansbergei assimilis
mariae
palawanus
pasia
pendleburyi
risi
semperi
williamsoni
| +++t++++++++
|
H++++++)]
(1) Dorsal stripe complete (+) / incomplete (—)
(2) Dorsal stripe connected (+) / not connected (—) with
mesothoracic half collar
(3) Yellow mark on frons separate (+) / one stripe (—)
(4) Anterior side of postclypeus with yellow patches (+)
/ dark (—)
(5) Yellow mark on vertex present (+) / absent (—)
A
We
_
| ++|HH++++]
VAN Tor: Malesian species of Leptogomphus
conspecificity is based on this character for several
species. Structural characters include the appen-
dages of the male and the vulvar scale of the female.
Also the tubercles on vertex and occiput are fre-
quently used, but these characters have to be con-
sidered carefully. I have seen several specimens
with very apparent differences in the tubercles on
each side of the head. Asahina (1986: 22) mentions
female specimens of L. gestroz Selys with an occip-
ital structure significantly different from the (allo)
type. This phenomenon was also observed by him
in Merogomphus parvus.
The distribution of the species was taken from
the following works: for India in the wider sense
(Laidlaw 1922, Fraser 1927, 1934), for Thailand
(and Burma) (Williamson 1907, Asahina 1986), for
Sumatra (Krüger 1899, Ris 1927, Lieftinck 1935,
1948), for Java (Lieftinck 1934), for Borneo (Laid-
law 1912, 1914, 1934, 1936, Lieftinck 1948) and for
the Philippine Islands (Needham & Gyger 1937,
Asahina 1968). Lieftinck (1954) is a most useful
summary of knowledge for the larger part of this
area.
Table 2 provides a data matrix for several impor-
tant characters used in the dichotomous key below.
I have examined all species myself.
1. Thorax with complete antehumeral stripe run-
ning close to the humeral suture (e.g. figs. 16-
17, 19-21)
Antehumeral stripe on thorax absent or
strongly reduced, usually to a small pale spot in
the posterior corner of the thorax above the
humeral suture (e.g. figs. 15, 18) 10
2. (1) Vertex with yellow marking; transversal
stripe on frons not interrupted by a longitudi-
nal black line inithe, middle; td. 3
(4) (5) (6) (7) (8) (9)
Ar — — — ap +
att + 4 im ? 24
= AL + = - ?
u I ie al n a
— — al — — ali
= i r = + ?
— _ r = SF Ar
EL di Da + ons (+)
— — r ? ? sr
= an 5A Al aif +
a = a = dE gE
+ + + a + ?
(6) Antehumeral stripe present (+) / absent (—) / re-
duced (r)
(7) Valve (female) far beyond segment 9 (+) / shorter (—)
(8) Tubercles on occiput in female present (+) / absent
=)
(9) Tubercles on abdominal segment 10 of male present
(+) / absent (—)
101
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
19 20
21
Figs. 15-21. - Thoracic pattern of left half of Leptogomphus species, dorsal view. - 15, L. pendleburyi, male holotype;
16, L. cf. pasia, male from Danum Valley; 17, L. williamsoni, female E. Borneo; 18, L. coomansi, male E. Borneo; 19,
L. lansbergei assimilis, male S. Sumatra; 20, L. gestroi, male Thailand, Doi Suthap; 21, L. #nclitus, female lectotype.
— Vertex black, without yellow or otherwise pale — Females ............... 7
markings; transversal stripe on frons inter-
rupted by a longitudinal black line in the middle
(CREfios ss MO Mio RE Tor. RER BRR. * 4
. (2) Characteristic markings on thorax with
very short dorsal stripe and large mesothoracic
half collar (fig. 17). Occiput black. Postclypeus
with latero-anterior corners pale coloured. Dis-
tabution#BorneoY.. nee ae.
ER EN AA ee L. williamsoni Laidlaw
Markings on dorsal side of thorax less charac-
teristic, with a mesothoracic half collar as in
most other congeneric species and a complete
dorsal stripe, which is not connected with the
mesothoracic half collar (fig. 20). Occiput with
a pale centre. Postclypeus brown, without pale
markings. Superior appendages of male, figs.
22-23. Distribution: Thailand, Burma. .......
OS calc ee er MNT L. gestrot Selys
Note. Redescription of male and female with illustra-
tions in Lieftinck (1960: 238-240, fig. 6).
MAO) IMAL ES sas aa EN Henk: 5
Note. The male of L. #nclitus (distribution Thailand)
is unknown. See fig. 21 for the thoracic markings of
the female. In all other species these markings agree
well between male and female.
102
. (4) Tip of superior appendage sharp with sub-
terminally a blunt ventral tubercle, thus in lat-
eral view bifid in appearance (figs. 34-35). Dis-
tributions Philippineiisiands aac, Mr mean
ER eee vai L. semperi (Selys)
Tip of superior appendage sharp, but without
subterminal ventral tubercle ……. 6
(5) Dorsal stripe (nearly) connected with me-
sothoracic half collar (fig. 5, 16); head pattern
as in fig. 8; superior appendages of male as in
figs: 32-334 Distribution: Borneo) .... ar
We tl eon oer we L. cf. pasia spec. nov.
Dorsal stripe and mesothoracic half collar se-
parated by a narrow black line; head pattern as
in fig. 10; superior appendages of male as in
figs. 30-31. Distribution: Java and Sumatra.
ite Soy N L. lansbergei (Selys)
Note. Specimens from Java belong to subspecies
lansbergei and specimens from Sumatra to subspecies
assimilis. According to Ris (1927: 28-29) the superior
appendages of both subspecies are different.
. (4) Valvula vulvae extremely long, extending to
the hind margin of segment 10 (figs. 3-4); oc-
ciput without tubercles (fig. 2); head pattern as
in fig. 1. Distribution: northern Borneo. .....
SIERRA GEE RSS REES L. pasta spec. nov.
VAN TOL: Malesian species of Leptogomphus
25 24
27
26 29
FTR
Figs. 22-37. - Leptogomphus superior appendages of male, for all species in dorsal and lateral view respectively. — 22-
23, L. gestroi from Thailand, Doi Suthap; 24-25. L. rist from Thailand, Nakhom Sithammarat province; 26-27,
L. coomansi from E. Borneo; 28-29, L. pendleburyi, holotype from northern Borneo; 30-31, L. lansbergei lansbergei from
W. Java; 32-33, L. cf. pasia from eastern Sabah, Danum Valley; 34-35, L. semperi from Basilan I.; 36-37, L. palawanus
from C. Palawan.
103
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
— Valvulae vulvae much shorter, extending not
further than the hind margin of segment 9
8. (7) Occiput with four slender spiny tubercles;
dorsal stripe connected with mesothoracic half
collar. Distribution: Philippine Islands. ......
DR RE tr NAR L. semperi (Selys)
Note. A female in the collection of the RMNH from
Mindanao keys out here. I consider this specimen
doubtfully conspecific with the male from Basilan I.,
which was compared with the type of L. semperi by
M. A. Lieftinck (for a detailed description of the male
specimen, see also Laidlaw 1936: 269).
— Occiput with ridges or smooth, but always with-
out spiny tubercles; dorsal stripe and mesotho-
racic half collar separated by a narrow black line
9. (8) Hind margin of occiput nearly straight or
with two roundish tubercles in the middle (size
of tubercles showing much variation). Labrum
black with a paired subrectangular pale spot (cf.
fig. 10). Last abdominal segments, fig. 12. Dis-
tributions avarandiSumattay nun.
Bee Metre A L. lansbergei (Selys)
Note. According to Lieftinck (1948, p. 245 and fig. 9)
specimens from Java (subspecies lansbergei tend to
have more conspicuous tubercles than specimens
from Sumatra (subspecies assimilis Krüger).
— Hind margin of occiput against both eyes with
an erect squarish tubercle (fig. 14). Labrum pale
(creamish yellow ?) with a narrow dark stripe
along the anterior margin and a very narrow
longitudinal dark line in the middle. Last ab-
dominal segments, fig. 13. Distribution: Thai-
landen er L. inclitus (Selys)
10. (1) Middle of postclypeus with a yellow patch;
antehumeral stripe absent or reduced to a small
spot in posterior part of synthorax above hu-
meral suture (fig. 18). Superior appendages of
male, figs. 26-27. Distribution: Borneo. ......
ERO ON. L. coomansı Laidlaw
— Postclypeus black or brownish black, without
yellow markings; antehumeral stripe reduced
to a larger or smaller pale spot in posterior part
of synthorax above humeral suture ...... 11
11. (10) Yellow mark on frons a single transversal
stripe (fig. 9); thoracic pattern, fig. 7; superior
appendages of male, figs. 28-29. Distribution:
Northern Borneo. .... L. pendleburyi Laidlaw
Note. Only known from male holotype.
— Transversal pale stripe on frons divided in the
middle, or at least with distinct dark triangular
emarginations in the middle ........... 12
12, MIG. a ean Ee 15
Note. The male of L. mariae is unknown.
TDA EN 14
13. Superior appendages in dorsal view curved in-
104
wards, tip bluntly shaped (figs. 24-25). Distri-
bution: Malaysias EP ea: L. risi Laidlaw
— Superior appendages in dorsal view straight,
tip bifid (figs. 36-37). Distribution: Palawan.
ENE MA L. palawanus Asahina
Note. Specimens from Borneo that key out here, may
belong to L. martae, of which the male is undescribed.
See Lieftinck (1948, p. 248) for an illustration of the
thoracic pattern of the female.
14. Occiput smooth, without tubercles. Distribu-
tion: Malayase o L. risi Laidlaw
— Occiput with conspicuous tubercles (Borneo
andiPhilippinesislands) We 15
15. Area behind lateral ocelli with a sharp tubercle,
directed posteriorly. Distribution: Palawan.
RS DO a Se L. palawanus Asahina
— Area behind the lateral ocelli with a semicircu-
lar transversal rim divided into two parts by a
central\deptessioni(Borneo) ME
SE Ee L. mariae Lieftinck
Note. This species is only known from the female
holotype. In other species with tubercles on the oc-
ciput, these tubercles may be absent or broken at one
or both sides of the head.
ACKNOWLEDGEMENTS
I am indebted to Mr. Stephen J. Brooks, British
Museum (Natural History) for his help during my
visits and for promptly sending type material under
his care, and to Dr. Erik J. van Nieukerken for his
comments upon a draft of this paper.
REFERENCES
Asahina, S., 1968. Records and notes on Philippine Odo-
nata. — Japanese Journal of Zoology 15 (4): 349-376,
figs. 1-58 + plate 1-2.
Asahina, S., 1970. Burmese Odonata collected by Dr. Ar-
thur Svihla with supplementary notes on Asiatic Ce-
riagrion species. — Japanese Journal of Zoology 16 (2):
99-126, figs. 1-68 + plate 1.
Asahina, S., 1986. A list of the Odonata recorded from
Thailand, Part XIV. Gomphidae-2. - Tombo 29: 7-53,
figs. 1-201.
Carle, F. L., 1986. The classification, phylogeny and bio-
geography of the Gomphidae (Anisoptera). I. Classi-
fication. - Odonatalogica 15 (3): 275-326, tables 1-2.
Fraser, F. C., 1927. Indian dragonflies. Part XXVI. - Jour-
nal of the Bombay Natural History Society 31(4): 882-
889, figs. 1-3.
Fraser, F. C., 1934. Odonata. Volume 2. - The Fauna of
British India including Ceylon and Burma: i-xxiii + 1-
398, figs. 1-119 + plate 1-4.
Krüger, L., 1899. Die Odonaten von Sumatra. IL. Theil.
Familie Aeschniden. — Stettiner entomologische
Zeitschrift 59 (1898): 267-331, figs.
Laidlaw, F. F, 1912. List of the Odonata taken on an
expedition to Mt. Batu Lawi together with descriptions
of supposed new species. — Journal of the Straits
Branch of the Royal Asiatic Society 63: 92-99, figs. 1-
DE
Laidlaw, F. F, 1914. Contributions to a study of the dra-
gonfly fauna of Borneo. — Part II. The Gomphinae and
Chlorogomphinae. — Proceedings of the Zoological
Society of London 1914: 51-63 + plate 1.
Laidlaw, F. F., 1922. A list of the dragonflies recorded from
the Indian Empire with special reference to the collec-
tion of the Indian Museum. Part V. The subfamily
Gomphinae. (with an appendix by F. C. Fraser). —
Records of the Indian Museum 24 (3): 367-426, figs.
EZRA
Laidlaw, F. F., 1931. A synonymic list of dragonflies of the
family Gomphidae (Odonata, Anisoptera) found in the
Oriental region. — Transactions of the Entomological
Society of London 78 (2): 171-197.
Laidlaw, F. F, 1933. Notes on Malaysian dragonflies (Odo-
nata), with descriptions of new species. — Bulletin of
the Raffles Museum, Singapore, Straits Settlements 7
(1932): 95-96, figs. 1-2b.
Laidlaw, F. E‚ 1934. A note of the dragonfly fauna (Odo-
nata) of Mount Kinabalu and of some other mountain
areas of Malaysia: with a description of some new or
little known species. - Journal of the Federated Malay
States Museums 17 (3): 549-561, figs. 1-4.
Laidlaw, F F, 1936. On some examples of the genus
Leptogomphus (Odon., Gomphidae). - Treubia 15 (3):
267-270, figs. 1-2.
Lieftinck, M. A., 1934. An annotated list of the Odonata
of Java, with notes on their distribution, habits and life-
history. - Treubia 14 (4): 377-462.
Lieftinck, M. A., 1935. A synopsis of the Odonata (Dra-
gonflies) of Sumatra. - Miscellanea Zoologica Suma-
trana 92/93: 1-23, figs. 1-2.
Lieftinck, M. A., 1948. Descriptions and records of South-
east Asiatic Odonata. - Treubia 19 (2): 221-278, figs.
1-19.
VAN Tor: Malestan species of Leptogomphus
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.
Lieftinck, M. A., 1960. On the identity of some little
known Southeast Asiatic Odonata in European mu-
seums described by E. de Selys Longchamps, with de-
scriptions of new species. - Memorie della Società
entomologica Italiana 38: 229-256, figs. 1-13.
Martin, R., 1904. Liste des Neuropteres de l’Indo-Chine.
- Mission Pavie Indochine 3: 204-221.
Needham, J. G. & M. K. Gyger, 1937. The Odonata of the
Philippines. - The Philippine Journal of Science 63
(1): 21-101, figs. 1-2 + plate 1-10.
Ris, F., 1927. Odonaten von Sumatra, gesammelt von
Edward Jacobson. - Zoologische Mededeelingen 10
(1): 1-49, figs. 1-24.
Selys Longchamps, E. de, 1878. Quatriemes additions au
synopsis des Gomphines. — Bulletin de l'Académie
royale de Belgique (2) 46: sep. 1-106.
Selys Longchamps, E. de, 1891. Viaggio di Leonardo Fea
in Birmania e regioni vicini. XXXII. Odonates. - An-
nali del Museo Civico di Storia Naturale di Genova
(serie 2a) 10: 433-518.
Williamson, E. B., 1907. The dragonflies (Odonata) of
Burma and Lower Siam. — II. Subfamilies Cordulegas-
terinae, Chlorogomphinae, and Gomphinae. - Pro-
ceedings of the United States National Museum 33
(1571): 267-317, figs. 1-39.
Received: 22 March 1990
Accepted: 24 March 1990
105
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
106
W. WITTMER
Naturbistorisches Museum, Basel
DIE FAMILIE MALACHIIDAE (COLEOPTERA)
AUF SULAWESI
43. Beitrag zur Kenntnis der indo-malaiischen Fauna
Wittmer, W., 1990. Die Familie Malachiidae (Coleoptera) auf Sulawesi. 43. Beitrag zur
Kenntnis der indo-malaiischen Fauna. - Tijdschrift voor Entomologie 133: 107-120, figs.
1-29. [ISSN 0040-7496]. Published 31 July 1990.
The species of the family Malachiidae occurring on Sulawesi are revised. Twenty-five species
are enumerated, of which nineteen are described as new to science. The genus
Luzonotroglops Pic, 1924 is synonymized with Falsolaius Pic, 1917, and the genus
Flabellapalochrus Pic, 1923 with Dromanthomorphus Pic, 1921. Carphurus tondanus
Champion, 1923 is synonymized with C. rufonotaticeps Pic, 1917.
Dr. W. Wittmer, Naturhistorisches Museum, CH-4001, Basel, Schweiz.
Key words. - Malachiidae, Sulawesi, new species.
FINLEITUNG
Von der Insel Sulawesi waren bisher nur 5 Arten
der Familie Malachiidae beschrieben worden und
zwar Carphurus celebensis Champion,
C. rufonotaticeps Pic, C. tondanus Champion,
Laius ribbei Pic und Dromanthomorphus cyaneus
Pic. Carphurus tondanus musste eingezogen wer-
den, weil diese Art Synonym mit C. rufonotaticeps
ist. Zwei weitere Arten: Carphurus cf. rubroannu-
latus Motschulsky und Latus cyaneus Guérin haben
eine weitere Verbreitung und werden heute zum
ersten Mal von Sulawesi gemeldet. Weitere 19 Ar-
ten stellten sich als neu für die Wissenschaft heraus.
Sie stammen von dem reichen Material, das ich
Herrn Dr. Peter Hammond, London, verdanke, das
im British Museum (Natural History) aufbewahrt
wird und vom “Project Wallace”, sponsored by the
Royal Entomological Society of London and the
Indonesian Institute of Science, Results of Project
Wallace No. 85 aufgesammelt wurde.
Den Londoner Koleopterologen, den Herren M.
J. D. Brendell, Dr. Peter Hammond, Dr. Christo-
pher Lyal und Dr. Nigel Stork, die auf Sulawesi so
erfolgreich tätig gewesen sind, gratuliere ich für die
schönen dort erzielten Resultate.
Des weiteren danke ich Herrn G. Hodebert, Pa-
ris, für die sorgfältig ausgeführten Zeichnungen
und Herrn Dr. Ch. O'Toole, Hope Museum, Oxford,
für die ausgeliehenen Typen.
Verwendete Abkürzungen: BM = British Mu-
seum (Natural History), London; MP = Muséum
de Paris, NHMB — Naturhistorisches Museum,
Basel.
SYSTEMATISCHER TEIL
Bestimmungstabelle für die auf Sulawesi
vorkommenden Gattungen
1. Die Flügeldecken bedecken das Abdomen voll-
kommen. Fühler anscheinend 10-gliedrig, das
Glied 2 steckt ganz im 1. Vordertarsen einfach,
oder das Tarsenglied 2 ist mit einem Kamm
versehen. „tonus Apt SERIE La er CRE 2
— Die Flügeldecken sind verkürzt und lassen
mehrere Tergite unbedeckt. Fühler 11-gliedrig,
das Glied 2 ist voll sichtbar. Glied 1 der Vorder-
tarsen mit einem Kamm (Carphurinae).... 3
. Fühlerglied 3 difform, meist mehr oder weniger
stark ausgehöhlt. Vordertarsen ohne Kamm
etant Seren 5 RN ER behe Laius Guérin
— Fiihler von Glied 3 an gekimmt. Glied 2 der
Vordertarsentmit Kamm RAEE
bes bapa sia wich cb: Dromanthomorphus Pic
3. Fliigeldecken einfach, ohne Auszeichnungen
N
— Flügeldecken gegen die Spitze mit einer mehr
oder weniger deutlichen Grube oder Eindruck,
mit vermehrten Haaren am Rand ..........
els enne e Paracarphurus Wittmer
4. Halsschild gegen die Basis schwach bis stark
verengt bis eingeschnürt, vor dem Basalrand
mässig bis stark quer eingedrückt. Fühler und
Korpegnormaljbehaactm nne: 3
— Halsschild vor der Basis schwach gerundet ver-
engt, Seiten davor fast parallel, ohne queren
Eindruck vor dem Basalrand. Körper und be-
sonders die Fiihler, meistens auch die Beine
107
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
RR E NR Telocarphurus Wittmer
5. Halsschild gegen die Basis schwach verengt,
vorderer Teil der Scheibe wenig bis kaum auf-
gewölbt, Eindruck vor dem Basalrand nur
schwach entwickelt. Rippen auf den Flügel-
cedken ENENMMMER NE ER eo
SE RIO IR RE e Carphurus Erichson
— Halsschild gegen die Basis stark eingeschniirt,
vorderer Teil der Scheibe stark gerundet aufge-
wölbt, Eindruck vor dem Basalrand breit. Rip-
pen auf den Flügeldecken fast immer vorhan-
ENTE: Falsolaius Pic
Carphurus Erichson, 1840
Bestimmungstabelle für die auf Sulawesi
vorkommenden Arten
1. Halsschild mit mehr oder weniger feinen Quer-
runzeln auf der Scheibe und an den
SEIEN. NN NR ION. 2
— Halsschild ohne Querrunzeln auf der Scheibe,
diese meistens glatt, schwach matt, oder punk-
GENE NN es chp cus RN 8
2. Halsschild und Flügeldecken schwarz, letztere
manchmal mit einer mehr oder weniger gros-
sen und deutlichen hellen (weisslichen) Auf-
hellung, oder vorwiegend weisslich .… 3
— Halsschild und Flügeldecken gelb oder hell-
bruno ere AMEN ER MELA 7
3. Flügeldecken einfarbig schwarz, oder mit mehr
oder weniger grossen und deutlichen hellen
(weisslichen) meistens seitlich gelegenen Auf-
hellungen, Naht immer dunkel .......... 4
— Flügeldecken fast ganz weisslich, nur an der
Basis schmal angedunkelt, Seiten und Spitzen
manchmal, jedoch noch schmäler als an der
Basis, angedunkelt ....... C. albipennis sp.n.
4. Fiihler kürzer, letzte 4 bis 8 Glieder angedun-
keltsbisischmarzins tn EED a 5
— Fühler länger (Abb. 2) einfarbig gelb, nur die
Spitze des letzten Gliedes ist dunkel ........
TR ae a C. pallidicornis sp.n.
5. Vorderschienen dunkel bis schwarz. Flügel-
decken glänzender, manchmal mit leichtem
violettem Schimmer, Punktierung etwas tiefer,
Elaaresschwarzigg ad. aaa mess 6
— Vorderschienen gelb. Flügeldecken weniger
glänzend, manchmal leicht matt, ohne Metall-
schimmer, Punkte etwas weniger tief, Haare
preis Aarden mer. C. dumogaensis sp.n.
6. Grössere Art: 6 mm (9). Fühler (Abb. 6) sehr
stark gezahnt (9). Flügeldecken schwarz mit
schwachem violettem Schimmer ...........
VAER: sine dona sia: C. celebensis Champion
— Kleinere Art: 4-5 mm (9). Fühler weniger
stark gezahnt (9). Flügeldecken schwarz, ohne
Metallschimmer oder heller (weisslicher) Auf-
108
122
155
HEID C. strigilatus sp.n.
. Der ganze Körper ist einfarbig gelb. Fühler
(Abb. 7) länger, alle Glieder länger als breit,
schwach gezahnt (8), einfarbig gelb oder die
letzten 1 bis 5 Glieder angedunkelt, selten ist
die Mittel- und Hinterbrust dunkel .........
SRI ee C. totopallidus sp.n.
Nur die Oberseite ist einfarbig gelb, auf der
Unterseite ist die Vorder- und Mittelbrust, mei-
stens auch die Hinterbrust schwarz, manchmal
auch das Abdomen, Basis der Mittel- und Hin-
terschenkel, seltener der Vorderschenkel dun-
kel. Fühler (Abb. 9) kürzer, einzelne Glieder so
lang wie breit oder breiter als lang (8), einfar-
big gelb oder ein paar der letzten Glieder sind
EEnansenaeieS Bo tare ee eee ee
UMILI AES ARE NEN C. partepallidus sp.n.
. Halsschild einfarbig rot oder orange bis gelb
heden Pe e O 9
Halsschild einfarbig schwarz oder zum grös-
Sn Tes AE RON O NNT 10
. Grosse Art: 5-6 mm. Halsschild rot, glatt, in
den Vorderecken befinden sich ein paar grobe
Punktes Fligeldeckenischwatz see eee
NE C. cf. rubroannulatus Motschulsky
Kleine Art: 2.5-3 mm. Halsschild orange bis
gelb, fein chagriniert, leicht matt. Fliigeldecken
gelblich, basale Hälfte bis ein Drittel dunkel
DALE AGREE IE C. basilimbatus sp.n.
. Halsschild einfarbig schwarz ........... 11
Halsschild an der Basis schmal gelblich ge-
STUER C. testaceolimbatus sp.n.
MIK PITCINfARDISISCAWA EZRA EN 12
Kopf rot, an den Schläfen und manchmal an der
Stirnbasis schmal schwarz. Flügeldecken
schwarz mit einem weisslichen mehr oder we-
niger ausgeprägten Langswisch ............
Nana mania ran C. brendelli sp.n.
Kopf zum grössten Teil glatt, vereinzelt mit
feinen Punkten besetzt. Halsschild ohne
PunktelanidentSel ent rt sun Dr or 13
Kopf mit ausgesprochen tiefen und groben
Punkten besetzt, besonders an der Basis, Zwi-
schenräume glatt. Halsschild an den Seiten
punktiert, Punkte an der Basis ineinander flies-
send. AN MOENS C. rufonotaticeps Pic
Kopf so breit wie der Halsschild, dieser mit
feinen Querrunzeln basal an den Seiten des
Quereindrucks. Punktierung der Fliigeldecken
wenig deutlich, fast erloschen .............
N ne Ken AEN OR TORE C. storki sp.n.
Kopf deutlich breiter als der Halsschild, dieser
ohne feine Querrunzeln basal an den Seiten des
Quereindrucks. Punktierung der Flügeldecken
dEUTLICHICODE PETER ARE C. lyali sp.n.
WITTMER: Malachiidae auf Sulawesi
1 mm
Abb. 1-6. Fühler von Carphurus. — 1, C. albipennis sp.n. 8; 2-3, C. pallidicornis sp.n. (2, 8. 3, 2); 4-5, C. dumogaensis
sp.n. (4, 8. 5, 2); 6, C. celebensis Champion 9.
Carphurus albipennis sp.n.
(Abb. 1)
Material. - Holotypus: &, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., lowland forest, ca. 200 m, 11.vii.1985
(BM). Paratypen: Gleicher Fundort wie Holotypus, ii, vi,
vii, x, xi und xii.1985 (14 BM, 12 NHMB); Fog 13, trays
81, 105, 106 (Plot A), 230/240 m, 11.vii.1985, (2 BM, 1
NHMB).
6. Kopf einfarbig schwarz, von der Mitte nach
vorne manchmal teilweise leicht, selten ganz aufge-
hellt; Maxillarpalpen gelb; Fühler gelb, die letzten
3 bis 5 Glieder mehr oder weniger dunkel; Hals-
schild, Schildchen und Abdomen schwarz; Fliigel-
decken weisslich, ringsum schmal, schwach ange-
dunkelt, manchmal nur angedeutet; Schenkel nur
an der Basis oder fast bis zu den Knien schwarz bis
dunkel, Schienen gelb oder ein wenig angedunkelt,
Tarsen gewöhnlich gelb, selten schwach angedun-
kelt.
Kopf mit den Augen breiter als der Halsschild,
Stirne leicht gewölbt, Querrunzeln an der Stirnbasis
nur angedeutet, restliche Oberfläche fast glatt, zer-
streute Haarpunkte sichtbar. Fühler (Abb. 1) die
Schultern ein wenig überragend, Glieder:4 bis 10
schwach stumpf gezahnt, einzelne Glieder ein we-
nig breiter als lang, 3 zur Spitze verbreitert. Hals-
schild kaum merklich breiter als lang, Seiten gerun-
det, gegen die Basis ein wenig stärker verengt als
nach vorne, Scheibe regelmässig gewölbt, vor der
Basis nicht quer eingedrückt und abgesetzt, Ober-
fläche dicht mit feinen Querrunzeln bedeckt. Flü-
geldecken nach hinten etwas erweitert, 3 bis 4 Ter-
gite unbedeckt lassend, fast glatt, zerstreut mit
ziemlich grossen, ganz flachen Haarpunkten be-
setzt. Behaarung des ganzen Körpers fein und greis,
nur auf dem Kopfe und Halsschild ein paar gröbere,
dunkle, längere Haare.
Länge: Knapp 3 mm.
Neben C. dumogaensis Wittmer zu stellen, der
nachfolgend beschrieben wird.
Carphurus pallidicornis sp.n.
(Abb. 2-3)
Material. - Holotypus: &, Indonesia, Sulawesi Utara:
Gng. Ambang F.R., near Kotamobagu, Gng. Muajat sum-
mit area, ca. 1780 m, 30.v.-2.vi.1985 (BM). Paratypen:
109
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
1 mm
aan
=
=
I
N
PESI
Abb. 7-12. Fühler von Carphurus. — 7-8, C. totopallidus sp.n. (7, 8. 8, 2); 9-10, C. partepallidus sp.n. (9, 8. 10, 2); 11,
C. basilimbatus sp.n. 8; 12, C. testaceolimbatus sp.n. 6.
Gleicher Fundort wie Holotypus, iv und xi.1985 (1 BM,
1 NHMB); Clarke Camp, lower montane forest, 1140 m,
x.1985 (1 BM, 1 NHMB); Sulawesi Tengah: Mt. Tambu-
sisi, 6500’, 01° 38 ’S-121° 23 ’E, 8-9.iv.1980, M.J.D. Bren-
dell (1 BM, 2 NHMB).
Kopf selten einfarbig schwarz und einer kleinen,
schlecht begrenzten hellen Makel zwischen den Au-
gen, oder der ganze Vorderkopf bis hinter die Füh-
lerwurzeln ist orangebraun; Fühler gelb, nur die
Spitze des letzten Gliedes ist mehr oder weniger
breit schwarz; Halsschild, Schildchen, Flügeldecken
und Unterseite schwarz, nur bei 1 Exemplar ist das
zweit- und drittletzte Tergit ein wenig aufgehellt;
alle Tibien und Tarsen gelb, bei den Mittel- und
Hintertibien manchmal ein wenig angedunkelt,
Schenkel schwarz, an der Spitze mehr oder weniger
breit gelb, bei den vorderen am breitesten gelb.
6. Kopf mit den stark vorstehenden Augen brei-
ter als der Halsschild, Stirne leicht gewölbt, einige
Querrunzeln an der Basis, davor grob punktiert, nur
in der Mitte eine kleinere fast punktfreie Stelle.
Fiihler (Abb. 2) ziemlich lang, ca. 2% mal so lang
wie der Halsschild, von Glied 3 an gezahnt, 3 ein
wenig länger als die folgenden. Halsschild kaum
merklich breiter als lang (22X21), Seiten kaum
gerundet, gegen die Basis ein wenig verengt,
Scheibe regelmässig gewölbt, vor der Basis nur ganz
schwach eingedrückt, Oberfläche dicht mit Quer-
runzeln bedeckt. Flügeldecken nach hinten leicht
erweitert, 2% bis 3 Tergite unbedeckt lassend, Ober-
fläche ziemlich dicht mit feinen manchmal etwas
erloschenen Punkten besetzt, Behaarung ziemlich
110
dicht, dunkel.
Q. Die Fühler (Abb. 3) sind kürzer als beim &,
weniger stark gezahnt, sonst mit dem @ überein-
stimmend.
Länge: 4-5 mm.
Ein wenig grösser als C. dumogaensis Wittmer,
mit diesem nahe verwandt, zu unterscheiden durch
die längeren, stärker gezahnten Fühler, die fast ein-
farbig gelb sind, ausgenommen die Spitze von Glied
11 und die durchwegs gelben Vorderschienen und
Tarsen.
Carphurus dumogaensis sp. n.
(Abb. 4-5)
Material. - Holotypus: @, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., G. Mogogonipa summit, 1008 m,
v.1985 (BM). Paratypen: Gleicher Fundort wie Holoty-
pus, v und xi.1985 (15 BM, 12 NHMB); Clarke Camp,
lower montane forest, 1140 m, iv, v und x.1985 (4 BM, 5
NHMB).
Schwarz bis schwarzbraun, Kopf auf der vorde-
ren Hälfte meistens mehr oder weniger rötlich bis
gelblich aufgehellt; Fühler gelb, die letzten paar
Glieder ein wenig angedunkelt oder ganz dunkel;
die letzten Tergite manchmal gelblich; Schenkel
meistens schwarz, oder an den Knien kurz gelb,
Tibien ein wenig aufgehellt, die vorderen meistens
ganz gelb, Tarsen gelb bis braun.
8. Kopf mit den Augen deutlich breiter als der
Halsschild, Stirne leicht gewölbt, Stirnbasis mit ein
paar deutlichen Querrunzeln, davor und neben den
Augen ziemlich grobe Punkte, die gegen die Mitte
und nach vorne rasch feiner werden. Fühler (Abb.
4) ungefähr so lang wie die Flügeldecken, Glieder
4 bis 10 ziemlich scharf gezahnt, 3 gegen die Spitze
verbreitert. Halsschild kaum merklich breiter als
lang, Seiten gerundet, gegen die Basis ein wenig
stärker verengt als nach vorne, Scheibe regelmässig
gewölbt, vor der Basis nicht quer eingedrückt und
abgesetzt, Oberfläche dicht mit Querrunzeln be-
deckt. Flügeldecken nach hinten leicht erweitert, 4
Tergite unbedeckt lassend, Oberfläche ziemlich
dicht mit feinen, etwas erloschenen Punkten be-
setzt, leicht matt wirkend. Behaarung wie bei
C. albipennis Wittmer.
®. Fühler (Abb. 5) ein wenig kürzer und weniger
breit gezahnt.
Länge: 3-3.8 mm.
Eine mit C. albipennis Wittmer nahe verwandte
Art, die sich durch etwas grössere Gestalt und die
dunklen, etwas stärker punktierten Flügeldecken
unterscheidet. Der Kopf ist bei dumogaensis im
Durchschnitt um ca. 15% breiter als der Halsschild
und misst im © 0.8 mm, bei albipennis ist er nur ca.
8-9% breiter als der Halsschild und misst im © 0.7
mm.
Carphurus celebensis Champion
(Abb. 6)
Carphurus celebensis Champion, 1923: 12.
Die Art wurde nach 1 9 aus Tondano, Celebes
beschrieben, der Holotypus befindet sich im Hope
Museum Oxford, ex Wallace. In dem vorliegenden
Material ist C. celebensis nicht vertreten. Die Art
ist grösser als alle anderen auf Sulawesi vorkom-
menden Arten mit quergerunzeltem Halsschild.
Für die Abb. 6 des Fühlers diente der Holotypus.
Der Fühler erinnert im Bau und in der Grösse an
C. pallidicornis sp.n. &, doch ist er bei celebensis
schwarz, nur die ersten Glieder sind hauptsächlich
auf der Unterseite aufgehellt. Der Kopf und Hals-
schild besitzen einen schwachen grünlichblauen
und die Flügeldecken einen blauvioletten Schim-
mer.
Carphurus strigilatus sp.n.
Material. — Holotypus: g, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., Banks of River Tumpah, i.1985 (BM).
Paratypen: Gleicher Fundort wie Holotypus, ii, iv, vi und
viii.1985 (16 BM, 12 NHMB); idem lowland forest, ca. 200
m, i, ii, iii, iv, v, vi, vii und viii.1985 (24 BM, 14 NHMB);
idem near Edward’s Camp, ca. 500 m, iv und x.1985 (5
BM); idem Fog 9, Tray 46, mixed crops, 1.iii.1985 (1 BM);
idem Hog’s Back Camp, lowland forest, 492 m, vii.1985 (1
BM).
Kopf vorwiegend schwarz, vorderer Teil bis über
die Fühlerwurzeln rötlichbraun, manchmal er-
streckt sich die helle Färbung bis zwischen die Au-
WITTMER: Malachiidae auf Sulawesi
gen; Fühler schwärzlich, erste 3 bis 5 Glieder gelb-
lich; Halsschild und Schildchen schwarz; Fliigel-
decken einfarbig schwarz oder schwärzlich, oder
jede Decke mit einer hellen Längsaufhellung, mehr
oder weniger deutlich, meistens mehr an den Seiten
gelegen, ohne sie zu berühren; Unterseite schwarz,
letzte 1 bis 3 Segmente oft gelbraun; Beine schwarz,
oft sind die Tarsen, meistens nur die vorderen,
aufgehellt.
Die übrigen Merkmale inklusive die Fühler stim-
men mit der Beschreibung von C. dumogaensis
überein, bis auf die Flügeldecken, die glänzender
sind mit gröberer Punktierung, Behaarung etwas
kräftiger, schwarz.
Länge: 4-4.3 mm.
Diese Art ist nahe mit C. dumogaensis Wittmer
verwandt, sie unterscheidet sich durch etwas grös-
sere Gestalt, die dunkeln Beine bei denen höchstens
die Vordertarsen gelblich sind, selten sind die Vor-
derschenkel an den Knien und die Vordertibien an
der Spitze kurz aufgehellt. Die Flügeldecken sind
von variabler Färbung, bei dumogaensis immer
dunkel, sie glänzen, haben tiefere Punkte und die
Behaarung ist kräftiger, dunkel, bei dumogaensis
greis.
Carphurus totopallidus sp.n.
(Abb. 7-8)
Material. — Holotypus: &, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., lowland forest, ca. 200/300 m,
viii.1985 (BM). Paratypen: Gleicher Fundort wie Holoty-
pus, ii, iii, iv, vii, viii, x, xi und xii.1985 (13 BM,8 NHMB);
idem Fog 1, trays 2, 5, 12, 87 (Plot A), 230 m, 5.11.1985 (3
BM, 1 NHMB); idem Fog 3, trays 6, 43, 47, 54, 55, 67, 85,
88, 95 (Plot B), 315 m, 8.ii.1985 (9 BM, 2 NHMB); idem
Fog 5, trays 8, 10, 17, 24, 27, 30, 31, 41, 43, 46, 47,52, 54,
72, 91, 101, 106 (Plot C), 400 m, 11.ii.1985 (21 BM, 2
NHMB); idem Fog 11. trays 23, 24, 36, 41, 43, 52, 60, 65,
67, 69, 71, 72, 74, 76, 82, 89, 93, 94, 95, 96 (Plot A), 230
m, 10.111.1985 (22 BM, 3 NHMB); idem Fog 13, trays 4,
9, 11, 12, 14, 22, 25, 26, 30, 34, 41, 50, 54, 68, 78, 79, 85,
86, 87, 88, 90, 93, 98, 99, 100, 103 (Plot A) (21 BM, 6
NHMB); idem Fog 15, trays 44, 60, 91 (PLot C) (3 BM);
idem Fog 26, trays 27, 74, 76, 77, 86, 102, 107, 112 (Plot
A) (10 BM).
Einfarbig blassgelb, ausgenommen die Fiihler,
bei denen die Glieder 5 oder 6 bis 11 dunkel sind,
seltener 8 bis 11, oder nur 11, selten sind die Fühler
einfarbig gelb; manchmal ist die Unterseite ange-
dunkelt.
8. Kopf mit den Augen breiter als der Halsschild,
fast so breit wie die Flügeldecken an den Schultern,
Stirne leicht gewölbt, Querrunzeln an der Stirnbasis
wenig ausgeprägt, davor wenige Punkte, die gegen
die Mitte der Augen erlöschen, von hier ab fast
glatt. Fühler (Abb. 7) ungefähr so lang wie die
Flügeldecken, Glieder länger als breit, gegen die
Spitze leicht verbreitert, 5 bis 10 unter sich unge-
fähr gleich lang. Halsschild ungefähr so lang wie
din
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
breit, Seiten gerundet, gegen die Basis kurz verengt,
Scheibe gewölbt, vor der Basis leicht quer einge-
drückt, dieser Teil ein wenig aufgewölbt; Oberflä-
che mit feinen Querrunzeln die von der Basis bis
kurz über die Mitte oder noch weiter nach vorne
sichtbar sind. Flügeldecken nach hinten leicht ver-
breitert, ein wenig klaffend, ca. 3 Tergite unbedeckt
lassend, Oberfläche bei einzelnen Individuen fast
glatt, höchstens Haarpunkte sichtbar, bei anderen
mit deutlicher, etwas erloschener Punktierung, Be-
haarung sehr fein, hell.
Q. Eigentlich nur durch das Fehlen des Kammes
am Glied 1 der Vordertarsen vom & verschieden,
denn die Fühler (Abb. 8) zeigen wenig Unter-
schiede, sie sind kaum merklich kürzer.
Länge: 2-3 mm, mit dem Abdomen, das bei den
Q oft stark vorsteht.
Neben C. testaceipennis Pic zu stellen, der je-
doch einen glatten, an der Basis ein wenig stärker
eingedrückten Halsschild besitzt. Die Fühlerglieder
sind bei testaceipennis zur Spitze weniger verbrei-
tert.
Carphurus partepallidus sp.n.
(Abb. 9-10)
Material. — Holotypus: &, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., lowland forest, malaise trap, ca. 200
m, ii.1985 (BM). Paratypen: Gleicher Fundort wie Holo-
Fog 13, trays 77, 82, 100 (Plot), 11.vii.1985 (3 BM); idem
Edward's Camp, lowland forest, malaise trap, ca. 664 m,
26.iv-28.v.1985, 1-16.x.1985 (2 BM); Sulawesi Tengah:
near Morowali, Ranu River area, ii und iii.1985 (3 BM).
Kopf, Halsschild und Schildchen orange; Fühler
gelborange, letzte 1 bis 5 Glieder mehr oder weni-
ger angedunkelt, selten einfarbig gelborange; Flü-
geldecken gelb; Mittel- und Hinterbrust dunkel,
Koxen, Trochanteren und Hinterschenkel manch-
mal bis über die Mitte dunkel, Mittelschenkel an der
Basis oft kurz dunkel, Vorderschenkel meistens ein-
farbig gelb, Tibien und Tarsen gelb; Hinterleib stel-
lenweise oft leicht angedunkelt, Rest gelblich, sel-
tener ganz dunkel.
8. Kopf mit den Augen breiter als der Halsschild,
Surne leicht gewölbt, Querrunzeln an der Stirnbasis
kaum erkennbar, diese Stelle ist mit mehreren
Punkten besetzt, die an den Seiten zahlreicher sind
und bis zu dem Hinterrand der Augen und nach
vorne bis zu den Fühlerwurzeln reichen, zwischen
den Augen fast glatt. Fühler (Abb. 9) verhältnis-
mässig kurz und kräftig, nicht ganz 2% mal so lang
wie der Halsschild, breit gezahnt, einzelne Glieder
ein wenig breiter als lang, 3 länger als 4. Halsschild
ein wenig breiter als lang (18X16), Seiten gerundet,
gegen die Basis ein wenig stärker verengt als nach
vorne; Oberfläche mit deutlichen Querrunzeln;
Scheibe gewölbt, an der Basis kaum angedeutet ein-
gedrückt. Flügeldecken nach hinten leicht verbrei-
112
tert, ca. 3 bis 4 Tergite unbedeckt lassend, Punkte
deutlich, etwas erloschen, ihr Abstand viel gròsser
als ihr Durchmesser, Behaarung sehr fein, weiss-
lich.
Q. Fühler (Abb. 10) ein wenig kürzer, weniger
breit gezahnt, sonstiger Bau und Skulptur wie beim
3.
Lange: ca. 3 mm.
Diese Art gehört in die Verwandtschaft von
C. albipennis Wittmer, sie unterscheidet sich
hauptsächlich durch die Färbung, vergleiche Be-
schreibung und Bestimmungstabelle.
Carphurus cf. rubroannulatus Motschulsky
Carphurus rubroannulatus Motschulsky, 1859: 64.
Material. — Indonesia, Sulawesi Utara: Dumoga Bone
N.P., lowland forest edge, ca. 200 m, malaise trap, ii und
iii.1985 (1 BM, 1 NHMB); Sulawesi Tengah: near Moro-
wali, Ranu River area, iii.1980, at light, M. J. D. Brendell
(1 BM, 1 NHMB).
Diese Art ist von Ceylon beschrieben und von
Indien, Tenasserim, Siam, Penang, Singapore, Java,
Sarawak, Laos, Tonkin, Philippinen, Palawan, Su-
matra, Molukken, Ceram, Buru, Borneo in der Lite-
ratur erwähnt. Aehnliche Arten sind aus Sumatra
(C. atromaculatus Pic), Burma (C. birmanicus Pic)
beschrieben. Aus Sulawesi liegen mir 4 9 vor, die
ich aufgrund ihrer Aehnlichkeit provisorisch als
rubroannulatus bestimme.
Carphurus basilimbatus sp.n.
(Abb. 11)
Material. — Holotypus: 6, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., Hog’s Camp, lowland forest, 492 m,
vii.1985 (BM). Paratypen: Gleicher Fundort wie Holoty-
pus, vii.1985 (1 NHMB); idem Fog 5, trays 1, 5, 46, 49,
52, 69, 70, 88, 100 (Plot C), 400 m, 11.11.1985 (6 BM, 3
NHMB); idem Fog 15, trays 81, 105, 106 (Plot C),
19.vii.1985 (2 BM, 1 NHMB); idem sites 10 and 11,
Tumpah Transect, 664 m, 19-25.11.1985 (1 BM).
Kopf und Halsschild einfarbig gelborange; Füh-
ler gelblich, letzte 3 bis 5 Glieder etwas angedun-
kelt; Schildchen und grösster Teil der Unterseite
mehr oder weniger aufgehellt; Flügeldecken gelb-
lich, Basis zu % bis % schwarz; Beine blassgelb.
8. Kopf mit den Augen breiter als der Halsschild,
Stirne leicht gewölbt, Stirne schmal, gegen den
Hinterrand der Augen breiter, ziemlich dicht punk-
tiert, zwischen den Augen glatt, hier stehen nur
ganz vereinzelte Punkte. Fühler (Abb. 11) ziemlich
lang, wenig mehr als 3 mal länger als der Hals-
schild, Glieder langlich, gegen die Spitze etwas ver-
breitert, 2 breiter als 3, ein wenig nach unten ver-
längert. Halsschild nur wenig breiter als lang
(16X15), Seiten gerundet, kurz vor der Basis ein
wenig verengt, Scheibe schwach gewölbt, vor der
basis schwach eingedriickt; Oberfläche fein punk-
tiert, stellenweise ein wenig gewirkt, dadurch leicht
matt wirkend. Flügeldecken nach hinten leicht er-
weitert, ca. 3 bis 4 Tergite unbedeckt lassend, fast
unpunktiert, leicht gewirkt, Behaarung fein, greis.
Letztes Tergit sehr kurz, breiter als lang.
Q. Die Fühler sind kaum merklich kürzer als
beim 4, Glied 2 ebenfalls nach unten verbreitert.
Länge: 2.5-3 mm.
Eine durch die Färbung der Flügeldecken sehr
charakteristische Art, dem C. semiflavus Wittmer
sehr ähnlich und die ausserdem durch das verdickte
Fühlerglied 2 auffällig ist, das bei semiflavus nicht
verdickt ist.
Carphurus testaceolimbatus sp.n.
(Abb. 12)
Material. — Holotypus: 6, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., Fog 3, tray 5 (Plot B), 315 m, 8.11.1985
(BM). Paratypen: idem Fog 12, tray 29 (Plot A), 280 m,
5.v.1985 (1 BM); idem Fog 26, tray 89 (Plot A), 230 m,
2.x11.1985 (1 NHMB).
8. Kopf, Schildchen und Abdomen schwarz-
braun; Fühler schwarz, erste 4 bis 5 Glieder gelb;
Halsschild schwarzbraun, basales Drittel bis Viertel
gelbbraun; Flügeldecken schwarzbraun, mit einem
gelblichweissen Querband vor der Mitte, ungefahr
so breit wie der Abstand zur Basis; Beine gelblich.
Kopf mit den Augen breiter als der Halsschild,
Stirne ziemlich flach, Schläfen runzlig gewirkt oder
mit Querrunzeln, die manchmal bis zur Stirnbasis
zu erkennen sind, Rest des Kopfes glatt mit ganz
vereinzelten Punkten. Fühler (Abb. 12) 3.5 mal so
lang wie der Halsschild, Glieder 3 bis 10 gezahnt,
unter sich ungefähr gleich lang. Halsschild unge-
fähr so lang wie breit, Seiten leicht gerundet, von
der Mitte zur Basis stärker verengt als nach vorne;
Scheibe gewölbt, vor dem Basalrand flach, jedoch
praktisch nicht eingedriickt; Oberfläche glatt, glän-
zend, wie der Kopf punktiert. Fliigeldecken fast
parallel, ca. 4 Tergite unbedeckt lassend, glatt glin-
zend, ein paar Haarpunkte kaum sichtbar, Behaa-
rung spärlich, schräg aufstehende kleine Borsten,
ausserdem 3 bis 4 längere Borsten auf jeder Decke.
Länge: 2-2.3 mm.
Deise Art unterscheidet sich von C. /yali sp.n.
durch den mit einem gelblichen Saum versehenen
Halsschild und dem gelblichweissen Querband auf
den Flügeldecken; die Fühler sind bei
testaceolimbatus ein wenig stärker gezahnt, die Flü-
geldecken praktisch glatt, bei C. /yali sp.n. deutlich
punktiert.
Carphurus brendelli sp.n.
(Abb. 13-14)
Material. — Holotypus: 8, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., Fog 11, tray 24 (Plot A), 10.111. 1985
WITTMER: Malachtidae auf Sulawesi
(BM). Paratypen: idem Fog 12, tray 112 (Plot A), 5.v.1985
(1 BM); idem Toraut alluv. forest, 1-2.11.1985 (1 NHMB);
idem Plot A, ca. 200 m, lowland forest, 6-13.111.1985 (1
BM); idem Edward’s Camp, lowland forest, 664 m, 26.iv-
7.vi.1985 und x.1985 (1 BM, 1 NHMB).
Kopf rot, Wangen und manchmal die Kopfbasis
schmal schwarz; Fühler schwarz, erste 4 bis 5 Glie-
der gelb; Halsschild, Schildchen, Unterseite und
Flügeldecken schwarz, letztere mit einem ver-
schwommenen bis deutlichen, weisslichen, mehr
oder weniger langen Längsflecken, der weder die
Naht noch die Seiten berührt; Beine gelb, beim
einzigen vorliegenden @ sind die Hinterschenkel
fast bis zu den Knien schwarz, die Hintertibien
angehaucht dunkel; bei den 9 sind oft auch die
Mittelschenkel fast bis oben, seltener die Vorder-
schenkel an der Basis kurz schwarz.
8. Kopf mit den Augen ein wenig breiter als der
Halsschild, Stirne leicht gewölbt, Schläfen an der
Basis mit Querrunzeln, an der Stirnbasis meistens
durch den Halsschildvorderrand verdeckt, davor
wenige gröbere Punkte, Rest des Kopfes praktisch
glatt. Fühler (Abb. 13) sehr lang, 4 mal so lang wie
der Halsschild, Glieder 5 bis 10 zur Spitze regelmäs-
sig leicht verbreitert, 4 ein wenig länger als 5, zur
Spitze stärker verbreitert als dieses, breit ausgeran-
det, 3 ein wenig kiirzer als 4, 2 nach unten leicht
gerundet erweitert, fast so breit wie 1. Halsschild
ein wenig langer als breit (24X22), Seiten nur
schwach gerundet, gegen die Basis verengt; Scheibe
in der Mitte ein wenig stärker gewölbt, vor dem
Basalrand deutlich quer eingedriickt; Oberfläche
glatt, ein paar Haarpunkte sichtbar. Flügeldecken
nach hinten leicht erweitert, ca. 5 Tergite unbedeckt
lassend, Punktierung spärlich, fast erloschen. Glied
1 der Vordertarsen verlängert und mit einem lan-
gen, an der Spitze gerundeten Kamm versehen.
©. Fühler (Abb. 14) einfach, wesentlich kürzer als
beim Ó.
Linge: 3.5-4.5 mm.
Färbung und Körperform wie bei C. rouyeri Pic,
verschieden durch den Bau des Fühlers beim Ó,
dessen Glied 4 breit ausgerandet ist.
Carphurus rufonotaticeps Pic
Carphurus rufonotaticeps Pic, 1917: 9.
Carphurus tondanus Champion, 1923: 35, syn.n.
Material. — Indonesia, Sulawesi Tengah: Ranu River
area, 27.i-20.iv..1980, sweeping clearings, M. J. D. Bren-
dell, BM 1980-280 (zahlreiche Exemplare BM, NHMB);
idem near Kolonodale, Gililana Village, 01°55’S-
121°22’E, 7-8.ii.1980, M. J. D. Brendell (6 BM); Sulawesi
Utara: Dumoga Bone N.P., Mogogompa, i, iii und x.1985
(14 BM, 14 NHMB).
Der Holotypus von Champion (Q) ist ca. 10%
grösser als die meisten mir vorliegenden Exem-
plare. Der Bau der Fiihler, Punktierung von Hals-
schild und Fliigeldecken stimmmt mit dem Paralec-
115
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
totypus ® von rufonotaticeps Pic überein, so dass
ich tondanus Champion als Synonym betrachte.
Carphurus storki sp.n.
(Abb. 15)
Material. — Holotypus: @, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., Rothamsted light trap, site 2, 220 m,
11.1985 (BM). Paratypen: idem Edward’s Camp, lowland
forest, 664 m, at light, x.1985 (1 NHMB); idem Fog 1, tray
56 (Plot A), 5.ii.1985 (1 BM).
Einfarbig schwarz, nur die Fiihlerglieder 1 bis 5
aufgehellt, 1 und 5 nur auf der Unterseite.
8. Kopf mit den Augen so breit wie der Hals-
schild, Stirne leicht gewölbt, an der Basis einige
=
Nn
III
/
|
5
feine Querrunzeln, davor ein paar Punkte, zwischen
den Augen und nach vorne glatt, spärliche Punkte.
Fühler (Abb. 15) knapp 3 mal so lang wie der
Halsschild, Glieder 3 bis 10 gegen die Spitze deut-
lich verbreitert, ein wenig länger als breit, 2
schwach nach unten erweitert, kaum merklich
schmäler als 3. Halsschild breiter als lang (19X17),
Seiten nach vorne schwach verengt, kurz vor der
Basis gegen diese verengt; Scheibe leicht gewölbt,
vor der Basis quer eingedrückt, an dieser Stelle
seitlich mit ein paar kurzen Querrunzeln, Rest
glatt, wenige Haarpunkte sichtbar. Flügeldecken
nach hinten ein wenig erweitert, 3 bis 3% Tergite
unbedeckt lassend, Punkte kaum angedeutet, erlo-
schen gewirkt, nicht glatt.
Q. Fühler um % kürzer als beim @.
Länge: 2.5-3 mm.
Die neue Art ist nahe mit C. brunneiventris (Pic)
verwandt. Obwohl die Picsche Art nach 1 © be-
schrieben wurde, ist C. storki leicht anhand des
Halsschildes zu unterscheiden, dessen Querein-
druck vor der Basis deutlich tiefer ist als bei
brunneiventris. Ausserdem ist er bei brunnet-
A mm
17
en
Abb. 13-17 — Fühler von Carphurus. 13-14, C. brendelli sp.n. (13, &. 14, Q); 15, C. storki sp.n. 8; 16, C. lyali sp.n. 6;
17, Fühler von Telocarphurus nebulosus sp.n. &.
114
ventris vollständig glatt, bei stork: zeigt er an den
Seiten ein paar gröbere, fast erloschene Punkte und
2 bis 3 deutliche kurze Querrunzeln an den Seiten
basal vor dem Quereindruck und eine grössere An-
zahl dahinter. Der Kopf zwischen den Augen ist bei
storki deutlicher punktiert als bei brunneiventris.
Carphurus lyali sp.n.
(Abb. 16)
Material. — Holotypus: @, Indonesia, Sulawesi Utara:
Gng. Ambang ER. near Kotamobagu, Fog 7, 1200 m,
18.11.1985 (BM).
8. Oberfläche und Abdomen einfarbig schwarz
bis schwarzbraun; Fühler schwarz, Glied gelb, 2
leicht aufgehellt. Beine braun.
Kopf mit den Augen viel breiter als der Hals-
schild, Stirne ziemlich flach, Stirnbasis gegen die
Schläfen und an denselben mit deutlichen Querrun-
zeln, diese dringen nach vorne neben den Augen bis
fast über die Fühlerwurzeln; zwischen den Augen
ganz wenige Punkte, vor der Stirnbasis gröbere und
zahlreichere Punkte. Fiihler (Abb. 16) fast 4 mal
länger als der Halsschild, Glieder 3 bis 10 schwach
gezahnt, 2 ungefähr so breit wie 3. Halsschild ein
wenig breiter als lang (13X12), Seiten wenig gerun-
det, kurz vor der Basis ein wenig verengt; Scheibe
gewölbt, vor der Basis kurz, flach eingedriickt;
Oberfläche glatt mit wenigen Haarpunkten. Flügel-
decken ca. 4 Tergite unbedeckt lassend, ziemlich
grob und dicht, teils etwas erloschen punktiert.
Länge: 2.7 mm.
Neben C. testaceolimbatus sp.n. zu stellen.
Telocarphurus Wittmer, 1939
Telocarphurus nebulosus sp.n.
(Abb. 17)
Material. — Holotypus: g, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., Fog 5, tray 9 (Plot C), 400 m,
11.11.1985 (BM).
8. Kopf, Halsschild, Schildchen und grösster Teil
der Unterseite schwarz; Fühler schwarz, 2 erste
Glieder gelb; Flügeldecken schwarz, basale °/; ver-
schwommen gelblichbraun; Beine braun bis
schwärzlich.
Kopf mit den Augen breiter als der Halsschild,
Stirnbasis sehr schmal und Schläfen gewirkt, Ober-
fläche glatt mit ein paar Punkten. Fühler (Abb. 17)
ca. 3% mal so lang wie der Halsschild, Glieder 3 bis
10 zur Spitze leicht verbreitert, unter sich ungefähr
von gleicher Länge, erst bei 10 deutlich kürzer als
3. Halsschild nur wenig breiter als lang (12X11),
Seiten nur schwach gerundet, Oberfläche glatt mit
wenigen Punkten, ausser je einer langen Borste in
den Basalecken ist die Scheibe mit einer Anzahl
WITTMER: Malachiidae auf Sulawesi
kürzerer Haare besetzt. Flügeldecken verkürzt, ca.
4 Tergite unbedeckt lassend, fast glatt, leicht durch-
sichtig, zerstreut, ungefihr wie der Halsschild be-
haart.
Linge: 2 mm.
Noch ein wenig kleiner und zarter gebaut als
T. strigilatus Wittmer, von diesem durch den glat-
ten Halsschild und die viel kürzeren Fühler leicht
zu unterscheiden.
Paracarphurus Wittmer, 1953
Paracarphurus sulawesiensis sp.n.
(Abb. 18)
Material. — Holotypus: 8, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., Clarke Camp, lower montane forest,
1140 m, iv.1985 (BM). Paratypen: Gleicher Fundort wie
Holotypus, v.1985 (1 BM); idem Edward's Camp, lower
montane forest, 664 m, x.1985 (1 NHMB).
8. Braun, letzte 4 bis 5 Fühlerglieder ganz
schwarz, manchmal auch die Spitzen ein paar wei-
terer Glieder angedunkelt, Hinterschenkel fast bis
zu den Knien und manchmal auch die Basis der
Mittelschenkel kurz schwarz; Unterseite schwarz,
letzte 3 Sternite orange, Tergite orange, ausgenom-
men das viertletzte, das schwarz ist.
Kopf mit den Augen kaum merklich breiter als
der Halsschild, Stirne nur schwach gewölbt, über
jeder Fühlerwurzel ein flacher Eindruck, Oberfläche
fast glatt, Stirnbasis mit ein paar Querrunzeln. Füh-
ler die Schulterbeulen ein wenig überragend, Glie-
der 3 bis 10 gezahnt, 3 ein wenig länger als 2.
Halsschild ein wenig länger als breit, Seiten vor der
Mitte am breitesten, hier gerundet, dann zur Basis
leicht verschmälert, Basis vor dem Basalrand leicht
aufgewölbt; Oberfläche glatt, feine zerstreute Haar-
punkte sichtbar. Fliigeldecken verkiirzt, etwas mehr
als doppelt so lang wie der Halsschild, jede Spitze
breit gerundet, fast in der Mitte, etwas vor der
Spitze befindet sich ein runder Eindruck mit leicht
erhöhtem Rand, vor dem Eindruck eine Anzahl
sehr lange weissliche, nach hinten gerichtete Haare
(Abb.18), die an der Spitze leicht verbreitert sind;
Punktierung mässig grob, Punkte teils zusammen-
fliessend, Zwischenräume glatt. Tarsalkamm kurz,
so lang wie das Tarsenglied 1.
Länge: 5 mm.
Die neue Art ist am nächsten mit P. neobrit-
tanicus Wittmer verwandt, sie hat ähnlich gerun-
dete Spitzen der Flügeldecken, die jedoch mit einem
grösseren Eindruck versehen sind, der weiter hin-
ten als bei neobrittanicus liegt. Bei neobrittanicus
fehlen die langen Haare hinter dem Eindruck, dafiir
befindet sich hier eine kurze breite Borste und zwi-
schen dem Eindruck und dem Hinterrand eine
grosse, fast runde, fast glatte Fläche, die bei
sulawesiensis fehlt.
115
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Falsolaius Pic, 1917
Luzonotroglops Pic, 1924 syn.n.
Der Vergleich des Holotypus von Luzonotrog-
lops carinatus Pic (spec. typ. von Luzonotroglops)
mit dem Holotypus von Falsolaius curtipennis Pic
(spec. typ. von Falsolaius), beide im MP, hat obige
Synonymie ergeben. Die beiden Arten F. carinatus
(Pic) und E curtipennis Pic bleiben bestehen. Alle
seit 1924 als Luzonotroglops beschriebenen Arten
sind zu Falsolaius zu stellen.
Die Gattung Falsolaius Pic ist nahe mit der Gat-
tung Neocarphurus Lea verwandt, die hauptsäch-
lich über Australien verbreitet ist, sie unterscheidet
sich von letzterer nur durch den einfachen Kopf,
dem Gruben oder sonstige Bildungen fehlen.
Falsolaius sulawesiensis sp.n.
(Abb. 19, 21)
Material. — Holotypus: 6, Indonesia, Sulawesi Utara:
Gng. Ambang F.R. near Kotamobagu, lower montane
forest, ca. 1750 m, 18-24.v.1985, yellow pan trap (BM).
mm
21 22
25
Paratypen: Gleicher Fundort wie Holotypus, 18-24.v.1985
(1 BM - @ in copula mit Holotypus — 2 NHMB); Gng.
Muajat, summit area, ca. 1780 m, vi.1985, malaise trap (1
BM).
Kopf und Halsschild braun, letzterer an der ein-
geschniirten Stelle bis zur Basis meistens mehr oder
weniger aufgehellt; Fühler braun, die letzten 2 oder
3 Glieder angedunkelt; Schildchen und Flügel-
decken braun, letztere oft mit einem schwach ange-
dunkelten Flecken auf jeder Schulterbeule, der stark
reduziert sein kann und nur noch neben dem Schild-
chen sichtbar ist, bei den Exemplaren mit gut ent-
wickelten Schulterflecken ist auch eine dunkle Mit-
telbinde vorhanden, die die Seiten und die Naht
nicht immer erreicht und je eine quere Makel an
den Spitzen; Abdomen dunkelbraun, letztes Tergit
heller; Beine braun, Schienen meistens ein wenig
angedunkelt, manchmal auch die Oberseite der
Schenkel.
8. Kopf mit den Augen breiter als der Halsschild,
Stirne schwach gewölbt, Basis kurz fein längsgestri-
chelt, Wangen ebenfalls gestrichelt, Rest des Kop-
fes gröstenteils chagriniert, zwischen den Augen
23
19 20 24
Abb. 18-25. — 18, Spitze der Flügeldecke von Paracarphurus sulawesiensis sp.n. &; 19-20, Fühler von Falsolaius (19,
E sulawesiensis sp.n. Ô, 20, E semicostatus sp.n. 8); 21-22, Letztes Tergit von Falsolaius (21, F. sulawesiensis sp.n. à;
22, E semicostatus sp.n. @); 23-25: Dromanthomorphus cyaneus Pic &. (23, Vordertibie von oben, daneben im Profil;
24, Mitteltibie; 25 letztes Tergit).
116
eine glattere Stelle, ein paar lange aufrechtstehende
Borsten vorhanden. Fühler (Abb. 19) fast 3 mal so
lang wie der Halsschild, Glieder 3 bis 10 zur Spitze
nur ganz wenig verbreitert, 4 länger als 3. Hals-
schild länger als breit (20X17), Seiten stark gerun-
det verengt, basales Drittel eingeschnürt und dann
zur Basis wieder ein wenig verbreitert, Seitenkante
vorne gegen den breiten und stark gewölbten Teil
am deutlichsten, gegen die Basis verschwindend,
der ganze eingeschnürte Teil ist körnig gewirkt,
matt, der vordere Teil glatt, vereinzelte Haarbor-
sten vorhanden. Flügeldecken nach hinten wenig
verbreitert, 4 bis 5 Tergite unbedeckt lassend,
Schultern stark vorstehend mit einer Längskante,
die nach den Schulterbeulen verschwindet, Spitze
jeder Decke schwach beulenartig aufgewölbt, Ober-
fläche körnig gewirkt, matt, mit zahlreicheren
schwarzen Borsten, besetzt als Kopf und Hals-
schild. Letztes Tergit (Abb. 21) ziemlich vorste-
hend, Spitze breit gerundet.
Q. Wie das 8 gebaut, sogar die Fühler sind kaum
merklich kürzer.
Die neue Art gehört zu den Arten mit glattem
Halsschild, wenigstens dem vorderen Teil und an-
gedeuteter Beule vor der Spitze der Flügeldecken.
Sie unterscheidet sich von F. unituberculatus (W itt-
mer) durch hellere Gestalt, den an der basalen,
verengten Stelle des Halsschilds, der fein gewirkt,
matt ist und die nur angedeutete beulenartige Ver-
dickung vor der Spitze der Flügeldecken, um nur die
wichtigsten Unterscheidungsmerkmale aufzuzäh-
len.
Falsolaius semicostatus sp.n.
(Abb. 20, 22)
Material. — Holotypus: 8, Indonesia, Sulawesi Utara:
Dumoga Bone N.P., Fog 5, trays 13, 14, 18, 26, 32, 54, 97
(Plot C), 11.11.1985 (BM). Paratypen: Gleicher Fundort
wie Holotypus, 11.11.1985 (4 BM, 3 NHMB); idem Fog 8,
trays 7, 11, 13, 15, 16, 18, 19, 20, 22, Mangrove, 3 km S.
of Labuanika, 0°52’N-23°57’E, 25.11.1985 (7 BM, 3
NHMB); idem Plot C, ca. 400 m, lowland forest, malaise
trap up trees, 18-25.1x.1985 (1 NHMB); idem Plot B, ca.
300 m, lowland forest, malaise trap up trees, 6-13.iii.1985
(1 BM); idem Edward's Camp, lowland forest, 664 m,
malaise trap, 26.iv-7.vi.1985 (1 BM).
Kopf dunkelbraun, vorderer Teil mehr oder we-
niger aufgehellt; Halsschild dunkelbraun, seltener
hellbraun, der eingeschnürte Teil an der Basis auf-
gehellt (gelblich); Fühler gelb, letzte 2 bis 5 Glieder
angedunkelt; Schildchen und Flügeldecken braun,
dunkelbraun oder seltener hellbraun, variabel auf-
gehellt, z.B. mit einer queren Aufhellung vor der
Mitte, die an der Naht unterbrochen sein kann und
einer breiteren dahinter, oder die hintere ist schmä-
ler als die vordere, bei dunkelbraunen Exemplaren
sind Aufhellungen kaum sichtbar oder nur ganz
schwach, manchmal nur hinten sichtbar; Abdomen
WITTMER: Malachiidae auf Sulawesi
dunkelbraun, die beiden letzten Tergite oder nur
das letzte kann hell sein; Beine dunkelbraun, oder
heller braun bis gelb, dann können die Schenkel,
manchmal auch die Schienen einen dunkieren
Längswisch aufweisen.
8. Kopf mit den Augen ein wenig breiter als der
Halsschild, Stirne leicht gewölbt, Oberfläche cha-
griniert (64X). Fühler (Abb. 20) fast doppelt so
lang wie der Halsschild, Glieder 3 bis 10 zur Spitze
nur ganz wenig verbreitert, 4 ein wenig länger als
3. Halsschild länger als breit (20X15), Seiten ge-
rundet verengt, basales Drittel eingeschnürt und
dann zur Basis wieder ein wenig verbreitert, Seiten-
kante auf dem vorderen Teil der Einschnürung kurz
oder nur angedeutet; Oberfläche am eingeschnür-
ten Teil fein chagriniert, Rest glatt. Flügeldecken
fast parallel, Schulterbeulen vorstehend, mit einer
Längskante, die ungefähr nach % verschwindet, vor
jeder Spitze ein Querwulst; Oberfläche fast glatt,
Behaarung fein gelblich nicht dicht, dazwischen
ganz vereinzelte, kurze aufstehende dunkle Bor-
sten. Letztes Tergit (Abb. 22) sehr kurz und breit,
Spitze ausgerandet.
Q. Fühler ein wenig kürzer als beim @, sonst wie
dieses gebaut.
Länge: 2-2.3 mm.
Nahe mit F sulawesiensis Wittmer verwandt,
Gestalt kleiner, die Seitenkanten an der Einschnü-
rung auf dem Halsschild sind weniger ausgeprägt,
die Querwulste an den Spitzen der Flügeldecken
sind grösser und das letzte Tergit ist viel kürzer,
Spitze ausgerandet.
Dromanthomorphus Pic, 1921
Flabellapalochrus Pic, 1923 syn. n.
Die Gattung Dromanthomorphus wurde von Pic
nach 1 ® aus Nord-Celebes, Toli-Toli, Nov.-Dez.
1895, H. Fruhstorfer, beschrieben, Holotypus im
MP. Im Material des BM fand sich 1 @, das zu
D. cyaneus Pic gehört. Pic hat bei der Beschreibung
der Gattung Flabellapalochrus nach 1 & übersehen,
dass er diese Gattung bereits nach 1 ® unter dem
Namen Dromanthomorphus beschrieben hatte, so-
dass erstere Gattung in Synonymie verfällt. Alle als
Flabellapalochrus beschriebenen Arten sind in die
Gattung Dromanthomorphus Pic zu stellen.
Dromanthomorphus cyaneus Pic
(Abb. 23-25)
Dromanthomorphus cyaneus Pic, 1921: 7.
Material. — Indonesia, Sulawesi Utara: Dumoga-Bone
N.P., Plot B. ca. 300 m, lowland forest, malaise trap,
ix.1985 (1 & BM).
Da das & noch nicht beschrieben wurde, folgt
hier dessen Beschreibung:
117
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
6. Kopf blau, Vorderkopf bis kurz über den Füh-
lerwurzeln, Oberlippen und Maxillarpalpen gelb;
Fühler gelb, die kammartigen Fortsätze der Glieder
4 bis 6 schwarz, 7 bis 11 ganz schwarz; Halsschild
und Schildchen blau; Flügeldecken weisslichgelb,
Basis und Spitzen ein wenig breiter, Seiten und
Naht schmal blau; Abdomen gelblich, letztes Tergit
braun, Spitze schwach, schmal angedunkelt; die 4
Vorderbeine gelb, Hinterbeine dunkel, an den
Knien leicht aufgehellt.
Kopf mit den Augen schmiler als der Halsschild
an der Basis, Stirne leicht, regelmässig gewölbt,
zwischen den Augen eine kleine Grube, Oberfläche
ziemlich grob punktiert. Fühler von Glied 4 an lang
gekämmt, 3 stark nach innen gerundet erweitert.
Halsschild breiter als lang, Seiten gerundet, ein
wenig nach vorne verengt, Scheibe gewölbt; Ober-
fläche unregelmässig, wenig dicht punktiert. Flü-
geldecken nach hinten leicht erweitert, fast ganz
erloschen punktiert, Behaarung doppelt, greis, kurz
auf der hellen Fläche, abstehende Borsten dunkel.
Letztes Tergit (Abb. 25) stärker sklerotisiert als die
nachfolgenden, lang, Spitze vorstehend, an den Sei-
ten ausgeschnitten. Vordertibien (Abb. 23) von
oben gesehen, vor der Spitze sehr schmal, wie zu-
sammengedriickt, im Profil gesehen (Abb. 23) nach
oben gerundet verbreitert; Mitteltibien (Abb. 24)
stark verbreitert, von oben gesehen ohne Aushöh-
lungen.
Länge: ca. 4.3 mm.
Dromanthomorphus ranuensis sp.n.
Material. — Holotypus: 8, Indonesia, Sulawesi Tengah:
near Morowali, Ranu River area, 27.i-20.iv.1980, Vert.
series 30 m actinic code, 13.111.1980, also at flight, M. J. D.
Brendell (BM); Paratypen: Gleicher Fundort wie Holoty-
pus (17 BM, 10 NHMB).
Kopf, Halsschild, Schildchen und Flügeldecken
schwarz mit mehr oder weniger deutlichem blauem
Metallschimmer, auf letzteren manchmal eine
längliche, schlecht begrenzte Aufhellung; Fühler
schwarz, erste 6 Glieder gelb, bei 4 bis 6 sind die
Kamme zunehmend angedunkelt; Beine schwarz,
Vordertarsen gelblich; Abdomen schwarz, die letz-
ten 2 bis 3 Sternite gelblich.
8. Kopf mit den halbkugelförmigen Augen brei-
ter als der Halsschild; Stirne schwach gewölbt;
Oberfläche dicht punktiert. Fühler um ca. ‘4 kürzer
als die Flügeldecken, von Glied 4 an in zunehmen-
der Lange gekämmt, 3 länglich, zur Spitze verbrei-
tert, diese stumpf, ein wenig gerundet. Halsschild
breiter als lang, Seiten gerundet, nach vorne etwas
verengt, Vorderecken etwas stärker gerundet als die
Basalecken; Oberfläche uneben, weniger deutlich
als der Kopf punktiert. Flügeldecken langgezogen,
nach hinten schwach verbreitert; Oberfläche teils
schwach runzlig, teils fast matt. Alle Beine ohne
Auszeichnungen.
118
Q. Augen normal, nicht vergrössert, Kopf mit
den Augen praktisch gleich breit wie der Hals-
schild. Fühler um ‘4 kürzer als beim @, stumpf
gezahnt.
Länge: ca. 3.3 mm.
Neben D. saigonensis (Pic) und D. impressitho-
rax (Pic) zu stellen, die beide ebenfalls einfache
Beine besitzen (8). Die neue Art ist durchwegs
grösser als die beiden erwähnten Arten von Pic, die
nur 2.5 und 2.8 mm messen.
Laius Guérin, 1830
Bestimmungstabelle für die auf Sulawesi
vorkommenden Arten
1. Kleine Arten, weniger als 3 mm messend; Flü-
geldecken mit 2 oder 4 kleinen weissen Flecken
— Grosse Art, 4 mm messend; Flügeldecken ein-
farbig blauviolett ........ L. cyaneus Guérin
2. Wangen einfach, ohne Gruben; Flügeldecken
mit 4aweissenthlecken) see CSO 3
— Wangen mit einer tiefen Grube, die sich bis zum
Hinterrand der Augen erstreckt; Flügeldecken
mit 2 weissen Flecken vor der Mitte. Fühler-
gliederail'-3; Abby I ER SARETE AE
aen miettes Har L. fossigerus sp.n.
3. Seiten des Halsschilds ungefähr in der Mitte
mit einem kräftigen Zahn ……. sanar: 4
— Seiten des Halsschilds einfach, ohne Zahn. Füh-
lerglieder 1-3, Abb. 27 ..... L. ranuensis sp.n.
4. Vorderkopf breit, über den Fühlerwurzeln
leicht abgesetzt aufstehend, Wangen flach;
Fühlerglied 3 kürzer, oben nicht längseinge-
drucke@Abbs26)i aa EEE L. ribbet Pic
— Vorderkopf nach vorne verschmälert, über den
Fühlerwurzeln nicht erhöht, Wangen ausge-
höhlt; Fühlerglied 3 länger, oben längseinge-
drucke(Albbs28) isan L. denticollis sp.n.
Laius fossigerus sp.n.
(Abb. 29)
Material. — Holotypus: 8, Indonesia, Sulawesi Tengah:
near Morawali, Ranu River area, 27.1-20.iv.1980, M. J. D.
Brendell (BM).
8. Kopf, Halsschild, Schildchen und Flügel-
decken schwarz, letztere mit 2 weissen ovalen, iso-
lierten Querflecken auf der vorderen Hälfte; Fiihler
schwarz, Glieder 2 bis 7 ganz, 8 basale Hälfte gelb;
Beine schwarz, alle Tarsen leicht aufgehellt.
Kopf mit den stark vorstehenden Augen breiter
als der Halsschild, Schläfen neben den Augen tief
ausgehöhlt, zwischen dem Hinterrand des Auges
und der Aushöhlung ragt eine schmale, wenig hohe
Leiste auf (von vorne gesehen gut sichtbar), Längs-
leiste zwischen den Augen am deutlichsten sicht-
bar, Wangen breit, flach; Oberfläche fein gewirkt,
matt. Fühler (Abb. 29) die Schulterbeulen nur we-
nig überragend, Glied 1 gegen die Spitze verbrei-
tert, 2 kaum sichtbar, 3 länglichoval, vor der Spitze
ein ovaler Eindruck, vor der Basis ein langer,
schmaler, schräg nach aussen gerichteter Fortsatz.
Halsschild ein wenig länger als breit, Seiten leicht
gerundet, Scheibe gewölbt, an der Basis quer einge-
drückt; Oberfläche fein chagriniert, matt. Flügel-
decken nach hinten leicht erweitert, ziemlich dicht
gewirkt, matt.
Länge: 2.5 mm.
Dies ist die einzige mir bekannte schwarze Lazus-
Art mit weissen Flecken auf den Flügeldecken, de-
ren Wangen tief ausgehöhlt sind.
Laius ranuensis n.sp.
(Abb. 27)
Material. — Holotypus: 8, Indonesia, Sulawesi Tengah:
near Morawali, Ranu River area, vert. series 20 m actinic,
14.11.1980, M. J. D. Brendell (BM).
6. Kopf schwärzlich, der vordere Teil der Wan-
gen und der Vorderstirne gelb; Fühler gelb bis gelb-
braun, die letzten 4 Glieder leicht angedunkelt;
Halsschild, Schildchen und Fliigeldecken schwärz-
lich, die letzteren mit 4 queren, weissen Flecken, die
vorderen erreichen die Seiten, die hinteren errei-
chen sie nicht ganz; Beine dunkel, die Spitze der
Vorderschenkel, die Vordertibien und alle Tarsen
gelb.
Kopf mit den Augen ungefähr so breit wie der
Halsschild, Stirne zuerst leicht gewölbt, dann bis
zur Vorderstirne fast glatt, Lingsleiste vorhanden,
26 Di
WITTMER: Malachtidae auf Sulawesi
an der Kopfbasis beginnend, nicht sehr deutlich
sichtbar, Wangen sehr breit; Oberfläche gewirkt,
matt. Fühler (Abb. 27) die Schulterbeulen ein wenig
überragend, Glied 1 keulenförmig, gegen die Spitze
nach aussen erweitert, 2 versteckt, 3 länglichoval,
an der Basis innen in eine kleine stumpfe, behaarte
Spitze ausgezogen, Oberfläche längseingedrückt.
Halsschild kaum merklich breiter als lang, Seiten
fast in gerader Linie gegen die Basis verengt;
Scheibe gerundet, vor der Basis schwach quer ein-
gedrückt, Oberfläche wie der Kopf gewirkt. Flügel-
decke nach hinten nur wenig erweitert, wie der
Halsschild gewirkt.
Länge: ca. 2.5 mm.
Neben L. denticollis Wittmer zu stellen, ver-
schieden durch die fehlenden Dorne an den Seiten
des Halsschilds und die anders geformten Fühler.
Laius ribbei Pic
(Abb. 26)
Laius ribbei Pic, 1910: 84.
Der Holotypus im MP trägt folgende Etikette: S.
Celebes, Pangie, C. Ribbe, 1882. Die Abb. 26 zeigt
die Fühlerglieder 1 bis 3.
Laius denticollis n.sp.
(Abb. 28)
Material. — Holotypus: @, Indonesia, Sulawesi Tengah:
near Morawali, Ranu River area, 27.1-20.1v.1980, M. J. D.
Brendell (BM). Paratypen: Gleicher Fundort wie Holoty-
pus (8 BM, 4 NHMB); Sulawesi Utara: Dumoga-Bone
N.P., lowland forest, ca. 200 m, 11.1985.
mm
28 29 1
Abb. 26-29. — Fühlerglieder 1 bis 3 von Latus @. — 26, L. ribbei Pic; 27, L. ranuensis sp.n.; 28, L. denticollis sp.n.; 29,
L. fossigerus sp.n.
119
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
6. Kopf, Halsschild, Schildchen und Flügel-
decken schwarz, letztere mit 4 weissen, isolierten,
fast runden Flecken, die 2 vorderen vor der Mitte,
die 2 hinteren hinter der Mitte; Fühler schwärzlich,
Glied 1 auf der Unterseite, 3 fast vollständig und die
folgenden 2 bis 4 auf der Innenseite bräunlich;
Beine schwärzlich, Tarsen ganz leicht aufgehellt.
Kopf mit den Augen kaum merklich schmäler als
der Halsschild an den seitlichen Zähnen, Stirne
gewölbt, eine feine Längsleiste beginnt an der
Stirnbasis und erstreckt sich bis ungefähr zur Mitte,
die Stirne ist vorne neben den Augen leicht gerun-
det aufgewölbt und dazwischen flach eingedrückt,
der grösste Teil der Wangen ist breit ausgehöhlt, so
dass die Vorderstirne sehr schmal ist; Oberfläche
gewirkt, matt. Fühler (Abb. 28) die Schultern ein
wenig überragend, Glied 1 keulenförmig gegen die
Spitze nach aussen erweitert, 2 fast ganz versteckt,
3 fast länglichoval, in der Mitte längseingedrückt,
an der Basis mit einem kurzen, leicht aufwärts ge-
richteten Zahn. Halsschild breiter als lang, Seiten
zur Basis verengt, jederseits fast in der Mitte ein
kräftiger Zahn, Oberfläche wie der Kopf gewirkt.
Flügeldecken nach hinten nur wenig erweitert, wie
der Halsschild gewirkt, matt.
120
Q. Wie das 8 gefärbt. Kopf einfach; Fühlerglied
1 zur Spitze verbreitert, jedoch nicht nach innen
erweitert, 3 so lang wie beim J, oben nicht einge-
drückt, Zahn an den Halsschildseiten kleiner.
Linge: 2.5-2.8 mm.
Neben L. celebensis Pic und L. dentatithorax Pic
zu stellen, die beide verschieden gebauten Kopf und
Fiihler besitzen.
LITERATUR
Champion, G. C., 1923. A revision of the Malayan and
Indian species of the melyrid subfamily Carphurinae
represented in the Hope Collection at Oxford and in
the British Museum in London (Coleoptera). - Annals
and Magazine of Natural History 9(12): 1-54.
Motschulsky, V. de, 1859. Etudes Entomologiques 8: 1-64.
Pic, M. 1910. Sur les “Laius” Guer. à quatre macules
blanches élytrales. - Echange 26: 83-84.
Pic, M., 1917. Descriptions abrégées diverses. - Mélanges
exotico-entomologiques 25: 1-24.
Pic. M., 1924. Coléoptères nouveaux des Philippines. —
Bulletin de la Société Entomologique de France: 1924:
230-231.
Received: 2 January 1190
Accepted: 30 April 1990
Tijdschrift voor Entomologie
Volume 133, no. I
| Editorial
Articles
3 A. Asquith & J. D. Lattin
Nabicula (Limnonabis) propinqua (Reuter) (Heteroptera: Nabidae): dimorphism,
phylogenetic relationships and biogeography.
17 A. Bedos & L. Deharveng
New species of Superodontella Stach (Collembola: Odontellidae) from Thailand.
27 ). Belle
Progomphus nigellus and Phyllocycla hamata, two new dragonflies from Brazil
(Odonata: Gomphidae).
3l K. A. Monk & R. K. Butlin
A biogeographic account of the grasshoppers (Orthoptera: Acridoidea) of
Sulawesi, Indonesia.
39 T. Pape
Two new species of Sarcophaga Meigen from Madeira and mainland Portugal
(Diptera: Sarcophagidae).
43 T. Pape
Revisionary notes on American Sarcophaginae (Diptera: Sarcophagidae).
75 V. Sruoga
Seven new species of Elachistidae (Lepidoptera) from the USSR.
85 Br. Theowald & P. Oosterbroek
Zur Zoogeographie der Westpaläarktischen Tipuliden. IX. Die Tipuliden des
Vorderen Orients.
97 J. van Tol
Key to the Malesian species of Leptogomphus Selys, with the description of a
new species from Sabah (Odonata, Gomphidae). Descriptions and records of
Malesian Odonata, |.
107 W. Wittmer
Die Familie Malachiidae (Coleoptera) auf Sulawesi. 43. Beitrag zur Kenntnis der
indo-malaiischen Fauna.
Book review
96 R. Johansson et al., The Nepticulidae and Opostegidae (Lepidoptera) of North
West Europa [R. T. A. Schouten]
© Nederlandse Entomologische Vereniging, Amsterdam
Published 31 July 1990 ISSN 0040-7496
Volume 133, no. 2, 1990 … ae ke i ISSN 0040-749
Tijdschrift
voor
Entomologie
A journal of systematic and evolutionary
entomology since 1858
. Published by the Nederlandse Entomologische Vereniging
Tijdschrift voor Entomologie
A journal of systematic and evolutionary entomology since 1858
Scope
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Co-editors
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RVARD
A a
GENUS CALATHUS (COLEOPTERA: CARABIDAE) ito, ERSITY
Aukema, B., 1990. Taxonomy, life history and distribution of three closely related species
of the genus Calathus (Coleoptera: Carabidae). — Tijdschrift voor Entomologie 133: 121-
141, figs. 1-18, tabs. 1-15 [ISSN 0040-7496]. Published 14 December 1990.
After cross-breeding experiments between Calathus melanocephalus, C. mollis mollis and
C. mollis cinctus and analysis of morphological data of these taxa, Calathus cinctus is
established as a good species. An identification key for the three species is given, main
morphological differences are depicted and data on their biotope, life history, distribution
and time trends are provided and discussed.
Berend Aukema, Pomona 66, 6708 CC Wageningen, The Netherlands.
Keywords. - Carabidae; Calathus, cinctus, melanocephalus, mollis, cross-breeding; identi-
fication key; life history; distribution maps; time trends.
In the past the status of Calathus melanocephalus
(Linnaeus, 1758) and C. mollis (Marsham, 1802) as
different species has been questioned many times,
mainly because of the frequent occurrence of mor-
phologically intermediate specimens. Schatzmayr
(1937), Lindroth (1943) and Wiebes-Rijks (1959)
treated them as distinct species, whereas Friede-
richs (1907) and Gersdorf (1937) could not find any
distinguishing character between both taxa at all.
Furthermore the existence of a morphologically
intermediate form cinctzs Motschulsky, 1850 (=
erythroderus Gemminger & Harold, 1868), estab-
lished as a subspecies of mollis by Lindroth (1945),
masks the status of these taxa.
This so-called melanocephalus group belongs to
Neocalathus, established as a subgenus of the genus
Calathus Bonelli by Ball and Negre (1972) with
Calathus melanocephalus as the type species. For
nomenclature in the melanocephalus group see Au-
kema (1990) and Aukema & Luff (1990).
The discovery at Wijster of populations of a cinc-
tus-like Calathus on abandoned agricultural fields in
the surroundings of the Biological Station of the
Agricultural University Wageningen (van Dijk
1978) offered a good opportunity for studying its
taxonomic relationship with both melanocephalus
and mollis, and its life history.
Moreover, in the present study cross-breeding
experiments between these three taxa were carried
out to elucidate their taxonomic status. Morpholog-
ical data (including biometrics) were used to sup-
port the resulting classification.
The taxonomic characters given in literature to
distinguish melanocephalus and mollis from each
other are:
l. The coloration of the pronotum in comparison
to the head (and elytra): clear rufous, contrast-
ing with the black head and elytra in melanoce-
phalus, and piceus to brown and not or little
contrasting in mollis. In the typical form of
melanocephalus and in mollis all appendages
are pale. However, a varying degree of mela-
nism may occur in melanocephalus, in wich the
pronotum is more or less infuscated, from pos-
sessing only a faint cloud at the centre to being
black with only narrowly translucent margins
(in that case also the appendages are melanistic,
at least tarsi brown, often also the main part of
legs and palpi, as well as middle antennal seg-
ments (Jeannel 1942, Lindroth 1974, 1986, Ve-
reshchagina 1985).
The shape and size of the right paramere: wi-
dened at the tip and hooked at the extreme apex
in melanocephalus and not widened and un-
armed or with a very small hook at the apex in
mollis (Lindroth 1974, 1986, Freude 1976, Ve-
reshchagina 1985). Differences in internal and
external morphology of the vesica are indicated
by Lindroth (1943) and Vereshchagina (1985).
3. The dimensions of the metepisterna: short in
melanocephalus (ca. 1.6 times as long as wide
in both brachypterous and macropterous spec-
imens), and longer in macropterous specimens
of mollis (ca. 1.8 times as long as wide) (Jeannel
1942, Lindroth 1974, Freude 1976, Vereshchag-
ina 1985).
N
1941
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
4. The shape of the pronotum: more slender, with
the greatest width usually before the middle
and clearly narrowed towards the base in mol-
lis, and less slender, with the greatest width in
or behind the middle and sides less convergent
in the basal half in melanocephalus (Freude
1976, Lindroth 1986, Vereshchagina 1985).
5. The shape of the elytra: longer oval (more
stretched), with lateral margin curved and
gradually tapering apically in mollis, and shor-
ter oval, with sides almost parallel and broadly
rounded apically in melanocephalus (Freude
1976, Lindroth 1986).
According to both Lindroth (1974, 1986) and
Freude (1976) the subspecies mollis cinctus (at the
time known as mollis erythroderus) can be distin-
guished from mollis mollis by the entirely rufous
pronotum, contrasting with the piceus to brown
head and elytra. Van Dijk (1978) furthermore men-
tions a difference in the colour of the sternites
between cinctus and melanocephalus. Material col-
lected in the field was studied to evaluate the use-
fulness of these characters for identification.
Fig. 1. Collecting sites of Calathus cinctus, C. melanoce-
phalus and C. mollis in The Netherlands. - 1, Drenthe
(Dwingeloo, Kraloo, Nuil); 2, Ameland (Buren, Nes); 3,
Texel ('t Horntje); 4, Oost-Flevoland (Ecological Re-
serve); 5, Amsterdam (Bijlmermeer), 6, Wassenaar
(Meijendel); 7, Schouwen (Haamstede).
122
Finally museum collections were revised in order
to get an impression of the distribution of the spe-
cies involved in The Netherlands. Due to misiden-
tification in the past, existing distribution maps of
the taxa concerned (Turin, Haeck & Hengeveld
1977) have to be replaced.
In this paper the taxonomic status of the three
taxa mentioned above is unravelled and the data
presented lead to the conclusion that Calathus cinc-
tus can be considered as a good species. Data on
habitat, life history and distribution of cinctus, me-
lanocephalus and mollis are given.
MATERIAL AND METHODS
Cross-breeding experiments
Single-pair crosses between all possible combina-
tions of the three taxa were carried out during
1978/1979 and 1979/1980 under approximately
outside conditions in an outdoor insectary. In
1978/1979 selected pairs were kept either in glass
50 km
jars on sieved peat litter (group 1) or in petri dishes
in ground peat litter (group 2). In the first case
newly emerged larvae were gathered from the peat
litter as described by van Dijk (1979a), whereas for
the second group eggs were separated from the peat
according the sieve-wash method of Mols et al.
(1981). Eggs were incubated at a constant temper-
ature of 19°C. Larvae were moved to glass tubes in
which they were reared individually in peat litter in
the outdoor rearing facility. During the winter
months (from the end of November until the end
of March) the larvae were moved to a climate room
at 5°C under short day (LD 10/14) conditions.
Single-pair crosses of males and females of the
same taxon served as control groups. Crosses and
origin of the beetles used are given in table 1.
Offspring of crosses performed by van Dijk
(1978) in 1977/1978 and specimens collected in the
field before the start of the reproduction period
(tenerals in the case of females; cinctus from Dwin-
geloo, melanocephalus from Kraloo, and mollis
from Voorne) were used as starting material (table
1). Furthermore, offspring of females inseminated
in the field (cznctzs from Nuil and Dwingeloo, and
melanocephalus from Oost-Flevoland) were reared
in order to get virgin females as starting material
for the next crossing programme.
In 1979/1980 crosses were carried out compara-
ble to those of group 2 of the preceding year. The
food, however, was changed from pieces of meal-
worm to small maggots. Again offspring of females
inseminated in the field (mollis from the West
Frisian Island of Ameland) was reared. Information
on these crosses is summarized in table 2. Collecting
sites are depicted in fig. 1.
Fig. 2. Measurements. BL: body length, BW: body width,
PL: length of pronotum, PW: greatest width of prono-
tum, EL: length of elytron, EW: greatest width of elytron,
WL: greatest length of wing, WW: greatest width of
wing, TIL: length of hind tibia, TAL: length of hind
tarsus.
AUKEMA: Three species of Calathus
Morphology
Material collected in the field was studied to
check the validity and usefulness of characters given
in literature for identification of the species con-
cerned. Samples of the following localities were
used:
Calathus cinctus: Drenthe: Dwingeloo (Noorden-
veld), Ruinen (Nuil); Friesland: Ameland (Buren,
Nes); N-Holland: Texel Ct Horntje); Zuid-Hol-
land: Wassenaar (Meijendel) and Zeeland:
Schouwen (Haamstede).
Calathus melanocephalus: Friesland: Ameland
(Buren, Nes); Drenthe: Dwingeloo (Noordenveld),
Ruinen (Nuil, Kraloo); Flevoland: Oost-Flevoland
(Oecologisch Reservaat); N-Holland: Texel Ct
Horntje), Amsterdam (Bijlmermeer).
Calathus mollis: Friesland: Ameland (Buren, Nes);
Z-Holland: Wassenaar (Meijendel) and Zeeland:
Schouwen (Haamstede).
Measurements were made with a Wild M5 ste-
reo-microscope provided with a calibrated ocular
micrometer. The following measurements were
taken (magnification between brackets):
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 1. Single-pair crosses between and within Calathus cinctus, C. melanocephalus and C. mollis in 1978/1979. cin:
cinctus; mel: melanocephalus, mol: mollis; lw: long winged; sw: short winged; N: number of crosses; NL: ibid, producing
larvae; L: number of larvae produced and reared; LR: number of larvae reared; B: number of beetles reared.
PARENTS
Species/Wing-morph Origin
Males Females Males Females N NL LAON
Group 1 mel sw cin lw Kraloo (28-6-1978) F;-offspring 1978!) 4 - - -
mel sw cin lw Kraloo (28-6-1978) Dwingeloo (19-7-1978, teneral) 1 - - =
mel sw cin sw Kraloo (28-6-1978) F -offspring 1978 5 = = =
cin lw mel sw F,-offspring 1978 Kraloo (28-6-1978, tenerals) 5 Lai -
cin sw mel sw F,-offspring 1978 Kraloo (28-6-1978, tenerals) 5 192 =
Group 2 cin lw mel sw Nuil (4-8-1978) F,-offspring 1978 6 - - -
cin lw mel sw F,-offspring 1978 F,-offspring 1978 6 - - -
cin sw mel sw Nuil (4-8-1978) F,-offspring 1978 2 - - -
cin sw mel sw F -offspring 1978 F -offspring 1978 7 = - -
mol lw mel lw Voorne (end July 1978)°) F,-offspring 1978 1 - - -
mol lw mel sw Voorne (end July 1978) F;-offspring 1978 5 = = =
cin lw mol lw Voorne (end July 1978) Voorne (end July 1978) 5 2 2Y -
mel sw mol lw Kraloo (4-8-1978) Voorne (end July 1978) 5 22 -
LR
Controls cin lw cin lw Parents 1977!) F -offspring 1978 6 DS)
cin sw cin lw F,-offspring 1978 Parents 1977 6 30 149759
cin lw no males Dwingeloo (23-8/22-9-1978) 16 16 734 471
cin lw no males Nuil (23-8/22-9-1978) 5 >) 259.198
cin lw cin sw F;-offspring 1978 Parents 1977 6 4 114 48
ein sw cin sw F -offspring 1978 F,-offspring 1978 6 6 295 1182
cin sw no males Dwingeloo (7-9-1978) 1 I 49 27
cin sw no males Nuil (23/31-8-1978) 4 3 147 86
mel sw mel sw Kraloo (30-8-1978) Kraloo (30-8-1978) 6 6 60 40
mel lw no males O-Flevoland (20-9-1978) 14 13 489 304
mel sw no males O-Flevoland (27-9-1978) gy CIO syle 236
'): parents and offspring of crosses carried out by Th. S. van Dijk
*): material collected by A. van Tiggele
ML1
Fig. 3. Measurements of met-episternum. ML1, ML2:
length; MW1, MW2: width. Scale 0.5 mm.
124
- length and width of the body (12): length between
tip of mandibles and apex of elytra and greatest
width over elytra (fig. 2: BL and BW).
- length and width of the pronotum (25): median
length and greatest width (fig. 2: PL and PW).
- length and width of the (right) elytron (12): EL
and EW in fig. 2. EW was calculated as half the
greatest body-width (0.5*BW).
- length and width of the (right) wing (12): greatest
length and greatest width (fig. 2: WL and WW).
- length and width of the (right) met-episternum
(50) as shown in fig. 3 (length: ML1, ML2; width:
MW1, MW2).
- length of the (right) hind tibia and of the (right)
hind tarsus (25) (fig. 2: TIL and TAL).
Elytra, wings, met-episterna and hind-legs were
removed from the body for proper measurement in
a flat plane. Wings were put on a slide in alcohol
and unfolded with a fine brush.
From these measurements ratios between length
and width of the body (BL/BW), width and length |
AUKEMA: Three species of Calathus
Table 2. Single-pair crosses between and within Calathus cinctus, C. melanocephalus and C. mollis (1979/1980). cin:
cinctus; mel: melanocephalus, mol: mollis; lw: long winged; sw: short winged; N: number of crosses; E: number of eggs
laid; NL: number of crosses producing larvae; L: number of larvae produced and reared; LR: number of larvae reared;
B: number of reared beetles.
PARENTS
Species/Wing-morph Origin
Males Females Males
mel lw cin lw Oost-Flevoland (11-10-1978)
mel sw cin lw F,-offspring 1979 (O-Flevoland)
mel sw cin lw F,-offspring 1979 (Kraloo)
mel lw cin sw Oost-Flevoland (11-10-1978)
mel sw cin sw F,-offspring 1979 (O-Flevoland)
mel sw cin sw F,-offspring 1979 (Kraloo)
mol lw cin lw Ameland (28-6-1979)
mol lw cin sw Ameland (28-6-1979)
cin lw mel lw F -offspring 1979
cin lw mel lw Ameland (28-6-1979)
cin Sw mel lw F -offspring 1979
cin sw mel lw Ameland (28-6-1979)
cin lw mel sw F,-offspring 1979
cin lw mel sw Ameland (28-6-1979)
cin lw mel sw F -offspring 1979
cin lw mel sw Ameland (28-6-1979)
cin sw mel sw F -offspring 1979
cin sw mel sw Ameland (28-6-1979)
cin Sw mel sw F,-offspring 1979
cin sw mel sw Ameland (28-6-1979)
mol lw mel lw Ameland (28-6-1979)
mol lw mel sw Ameland (28-6-1979)
mol lw mel sw Ameland (28-6-1979)
Control groups
mel lw mel lw F -offspring 1979
mel sw mel lw F,-offspring 1979 (O-Flevoland)
mel sw mel lw F;-offspring 1979 (Kraloo)
mel lw mel sw F,-offspring 1979
mel sw mel sw F -offspring 1979 (O-Flevoland)
mel sw mel sw F;-offspring 1979 (Kraloo)
mel lw mel sw F -offspring 1979
mel sw mel sw F -offspring 1979 (O-Flevoland)
mel sw mel sw F -offspring 1979 (Kraloo)
cin lw cin lw F -offspring 1979
cin lw cin lw Ameland (28-6-1979)
cin sw cin lw F -offspring 1979
cin sw cin lw Ameland (28-6-1979)
cin lw cin sw F -offspring 1979
cin lw cin sw Ameland (28-6-1979)
mol lw no males
Females = NIEREN TE
F,-offspring 1979 4 422 5) 4168 -
F,-offspring 1979 4 591 - - -
F -offspring 1979 4 320 = È _
F,-offspring 1979 4 91 - - -
F,-offspring 1979 4 219 = = È
F,-offspring 1979 AN u Bs à
F,-offspring 1979 4 184 = 2 =
F -offspring 1979 4 254 = = =
Oost-Flevoland (25-10-1978)
Oost-Flevoland (25-10-1978)
Oost-Flevoland (25-10-1978)
Oost-Flevoland (25-10-1978)
F,-offspring 1979 (O-Flevoland)
F,-offspring 1979 (O-Flevoland)
NNNNNNNNNNNN
WN
O0
F,-offspring 1979 (Kraloo) 218 DE -
F,-offspring 1979 (Kraloo) 14 - = -
F,-offspring 1979 (O-Flevoland) 15 = = -
F,-offspring 1979 (O-Flevoland) 50 2 -
F,-offspring 1979 (Kraloo) 87 -
F,-offspring 1979 (Kraloo) 4 - -
Oost-Flevoland (25-10-1978) 4 181 25 -
F,-offspring 1979 (O-Flevoland) 4 57 - -
F,-offspring 1979 (Kraloo) 459 = = =
LR
F,-offspring 1979 4 1173 4 238 85
F,-offspring 1979 4 1084 4 240 135
F,-offspring 1979 4 1279 A Zilk 9)
F,-offspring 1979 (O-Flevoland) 4 1113 4239 92:
F,-offspring 1979 (O-Flevoland) 4 959 4 240 130
F,-offspring 1979 (O-Flevoland) 4 665 4 224 136
F,-offspring 1979 (Kraloo) 4 682 4 160 96
F,-offspring 1979 (Kraloo) 4 1079 4 240 132
F;-offspring 1979 (Kraloo) 4 920 4 240 112
F,-offspring 1979 2 468 21207243
F,-offspring 1979 157232 17607520
F;-offspring 1979 2 639 2585) el
F -offspring 1979 1 509 IGOR
F,-offspring 1979 2 496 271167168
F,-offspring 1979 152216 122607729
Ameland (28-6-1979) 28 573 9 309 194
of the pronotum (PW/PL), pronotum width and
body width (PW/BW), length and width of the
elytra (EL/EW), length of the elytra and body
length (EL/BL), length and width of the wings
(WL/WW), wing-length and length of the elytra
(WL/EL), length and width of the met-episterna
(ML2/MW2), and the ratio between the lengths of
the hind tarsus and hind tibia (TAL/TIL) were
estimated.
Special attention was paid to the coloration of the
beetles, the size and shape of the right paramere
and to the number of dorsal punctures in the third
elytral interval.
125
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Habitat and life history
Details of the habitat and the life history of the
species were derived from the analysis of field data
(mixed populations of cinctus and melanocephalus
from Dwingeloo, Ruinen and the West Frisian Is-
land of Texel and populations of cinctus and mollis
from the West Frisian Island of Ameland), from
literature (populations of melanocephalus studied
by Vliim & van Dijk (1967), Vlijm et al. (1968) and
van Dijk (1972, 1973)) and from breeding experi-
ments with the three species under outside condi-
tions carried out in 1981/1982 and 1982/1985.
Distribution
Calathus-material of this group from Dutch col-
lections was revised to be able to get an impression
of the distribution of the species in The Nether-
lands. Material from the following collections is
included:
Museums and institutions: Zoölogisch Museum,
Amsterdam; Rijksmuseum van Natuurlijke Histo-
rie, Leiden; Vakgroep Entomologie, Landbouwuni-
versiteit, Wageningen; Biologisch Station, Land-
bouwuniversiteit, Wijster.
Private collections: K. den Bieman, Bennekom;
E. H. M. Bouvy, Beek; H. T. Edzes, Nijmegen; T.
van Gijzen, Arnhem; Th. Heijerman, Wageningen;
S. van Heijnsbergen, Naarden; Y. Jongema, Wage-
ningen and A. P. J. A. Teunissen, Vlijmen.
Additional material was supplied by K. Alders,
Arnhem; M. A. Baars, De Koog, Texel (Loopkever-
werkgroep Texel); Th. S. van Dijk, Wijster; P. J. M.
Mols, Heteren and H. Turin, Renkum.
Time trends in the occurrence of the species
(i. e. numbers of occupied 10 X 10 km squares of the
distribution maps per decade since 1870) were es-
timated according to the method of Turin & den
Boer (1988).
RESULTS
Cross-breeding experiments
In total 123 single-pair crosses were carried out:
59 between cinctus and melanocephalus, 47 be-
tween cinctus and mollis and 17 between melanoce-
phalus and mollis (tables 1 and 2). None of these
crosses produced adult offspring although in 5 (8.5
%), 3 (6.4 %) and 3 (17.6 %) cases respectively
larvae were produced, which, however, never
reached the third larval stage.
Table 3. Measurements (in mm) of short winged males of Calathus cinctus and C. melanocephalus from Texel. N:
number of measurements.
cinctus melanocephalus
MEASUREMENT N Mean 95% cl. N Mean 95% cl.
Body Length (BL) 20 6.79 6.59-6.98 20 6.66 6.51-6.82
Width (BW) 20 2.71 2.64-2.77 20 2.67 2.61-2.73
BL/BW 20 DL 2.47-2.55 20 2.50 2.45-2.55
Pronotum Length (PL) 20 1.64 1.61-1.68 20 1.68 1.65-1.72
Width (PW) 20 199) 1.95-2.04 20 2 2.06-2.15
PW/PL 20 1.21 1.20-1.23 20 1825 1.23-1.27
PW/BW 20 0.74 0.73-0.74 20 0.79 0.78-0.80
Elytron Length (EL) 20 8 4.07-4.29 20 4.09 4.00-4.18
Width (EW) 20 195 1.32-1.38 20 1.99) 1.30-1.36
EL/EW 20 3.09 3.05-3.12 20 3.07 3.03-3.11
EL/BL 0 0.62 0.61-0.62 20 0.61 0.60-0.63
Wing Length (WL) 20 1.93 1.85-2.01 20 1.93 1.86-2.00
Width (WW) 20 0.48 0.45-0.51 20 0.51 0.47-0.54
WL/ WW 20 4.05 3.86-4.24 20 3.86 3.65-4.07
WL/EL 20 46 0.45-0.47 20 0.47 0.46-0.49
Met-episternum Length (ML2)') 20 0.76 0.74-0.78 20 0.70 0.68-0.73
Width (MW2)!) 20 0.57 0.55-0.59 20 0.57 0.55-0.58
ML2/MW2 (ME2) 20 1.34 1.32-1.36 20 1.24 1.22-1.27
Hindleg Length tarsus (TAL) 20 2.07 2.01-2.13 20 2.02 1.98-2.06
Length tibia (TIL) 20 2.02 1.98-2.07 20 1.98 1.94-2.01
AE DIE 20 1.02 1.01-1.03 20 1.02 1.01-1.04
Dseertie 8;
All control crosses (42 of melanocephalus and 30
of cinctus) crosses that produced larvae (respec-
tively 100 % and 82 % for both years together), also
produced adult offspring (tables 1 and 2). In
1979/1980 all crosses of both species produced
offspring (table 2). In the case of mollis not suffi-
cient material to carry out control crosses in the
same way was available. However, of 28 females
collected in the field (Ameland, Buren, 28.vi. 1979)
and kept without males under the same conditions
as single-pair crosses, 9 produced larvae and off-
spring (table 2: 194 beetles from 309 larvae). The
other 19 females (ten of which were tenerals at the
time of capture) produced no or only a few unfer-
tilized eggs, and were considered not to have been
inseminated at all. Nevertheless it is clear from
these results that also in the case of mollis the
failure of single-pair crosses with cinctus and me-
lanocephalus cannot be attributed to the rearing
conditions used.
In 1979/1980 egg production was also estimated.
It appeared that, although unfertilized females may
produce a considerable number of eggs, the egg
production of fertilized females generally is much
higher (table 2). It is concluded that cinctus, mela-
nocephalus and mollis show mutual reproductive
isolation, and, therefore, have to be considered as
distinct species.
AUKEMA: Three species of Calathus
Morphology
Biometric data, especially the ratio length/ width
of the met-episterna, have been used in attempts to
separate the taxa of this group of closely related
species (Gersdorf 1937, Wiebes-Rijks 1959). The
establishment of cinctus as a valid species as the
result of the breeding program necessitated a new
attempt to find morphological differences between
the three species. Concerning biometrics, however,
two points have to be taken into account. Firstly
general differences both between sexes (on average
females are larger than males) and between wing-
morphs (on average long winged specimens are
larger than short winged specimens and differently
shaped) exist in these species. Secondly, the way
measurements are made should be described (and
figured) in detail, which apparently is not normal
practice. Lindroth (1974) and Freude (1976), for
instance, use the ratio length/width of the met-
episterna without indicating how to measure it, and
do not point out the differences between sexes and
between wing-morphs in a proper way.
Tables 3-6 list measurements and ratio's of both
males and females of short as well as long winged
Calathus cinctus and C. melanocephalus collected
on Texel by M. A. Baars and of long winged C.
mollis collected in the coastal dunes near Haam-
Table 4. Measurements (in mm) of short winged females of Calathus cinctus and C. melanocephalus from Texel. N:
number of measurements.
cinctus melanocephalus
MEASUREMENT N Mean 95% cl. N Mean 95% cl.
Body Length (BL) 20 1559 7.23-7.56 20 732 ~7.18-7.46
Width (BW) 20 3.01 2.94-3.09 20 2.95 2.91-3.00
BL/BW 20 2.46 2.42-2.50 20 2.48 2.44-2.52
Pronotum Length (PL) 20 177 1.73-1.81 20 1.79 1.76-1.82
Width (PW) 20 2,19) 2.13-2.24 20 2.28 2.24-2.32
PW/PL 20 1.23 16222125 20 1.27 1.26-1.28
PW/BW 20 0.73 0.72-0.74 20 0.77 0.76-0.78
Elytron Length (EL) 20 4.54 4.44-4.64 20 4.45 4.36-4.53
Width (EW) 20 Lil 1.47-1.54 20 1.48 1.46-1.50
EL/EW 20 3.02 2.98-3.05 20 3.01 2.97-3.04
EL/BL 20 0.61 0.61-0.62 20 0.61 0.60-0.62
Wing Length (WL) 20 2.14 2.08-2.20 20 2.06 1.99-2.13
Width (WW) 20 0.60 0.57-0.62 20 0.60 0.55-0.64
WL/WW 20 3.61 3.48-3.75 20 3.49 3.32-3.65
WL/EL 20 0.47 0.46-0.48 20 0.46 0.45-0.48
Met-episternum Length (ML2) 20 0.83 0.80-0.85 20 0.76 0.74-0.77
Width (MW2) 20 0.64 0.62-0.65 20 0.62 0.61-0.64
ML2/MW2 (ME2) 20 1.30 1.27-1.33 20 1.22 1.20-1.24
Hindleg Length tarsus (TAL) 20 2.08 2.03-2.12 20 197 1.92-2.01
Length tibia (TIL) 20 2.07 2.03-2.12 20 2.02 1.98-2.06
TAL/TIL 20 1.00 0.99-1.02 20 0.97 0.96-0.98
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 5. Measurements (in mm) of long winged males of Calathus cinctus and C. melanocephalus from Texel and C.
mollis from Haamstede. N: number of measurements.
cinctus melanocephalus mollis
MEASUREMENT N Mean 95% cl. N Mean 95% cl. N Mean 95% cll.
Body Length (BL) IRR 52-7057 TMD OSB) U ee TEN
Width (BW) 1 CAMP 55-2573 0 ar 2:39, aD 70) 29) Do
BL/BW AZ, IED 2.52 244-260 20 2.63 2.61-2.66
Pronotum Length (PL) 160 AE Co LOL TO STE 20,169: 1:61.69
Width (PW) U Oe ESS 15997 D'LA AT SI 205. 2205) 2.002209,
PW/PL This ZINN TENS Zed LMD TE ALD)
PW/BW RI OT. 072220, 74 7) 0.79 0.78-0.79 20 0.73 0.73-0.74
Elytron Length (EL) 1197 212705 3.98-4257 432 419-445 20 438 427-449
Width (EW) br SZ AEN 7 PASS SEAT 20771995 31362142
EL/EW LIN 31030, 31030 EON 20 ESE
EL/BL Ue OHG OEE 97 0.60 0.58-0.62 20 0.60 0.59-0.60
Wing Length (WL) 1.2629 SIL 7.10 686-785 20 6.61 6.39-6.83
Width (WW) I 12.36 223250 7 2.68 263-274 20 2.46 237-254
WL/WW Il m 2.097 Pos DOSE 258-25) 220i 2:69 ae 66-27/8,
WL/EL u 147157007 SA 16216720 IODS
Met-episternum Length (ML2) U OT 07420797 0.77 0.73-0.81 20 082 0.80-0.84
Width (MW2) STe LOOD 0.60 0.58-0.62 20 058 0.56-0.59
ML2/MW?2 (ME2) IE > E o 7 1.28 121-135 20 143 140-145
Hindleg Length tarsus (TAL) UO OEE 6 DEF NEP WA DO ZD ZLD DS)
Length tibial (TIL) 110 221:96.591.902203556 2.13 2.04-2.22° 20. 2:09), 2.04-2.15
TAL/TIL 11 102 101-104 6 102 1.00-1.05 20 1.05 1.04-1.06
Table 6. Measurements (in mm) of long winged females: Calathus cinctus and C. melanocephalus from Texel and C.
mollis from Haamstede. N: number of measure-ments.
cinctus melanocephalus mollis
MEASUREMENT N Mean 9%cl N Mean 95% cl. N Mean 95% cl.
Body Length (BL) 11 927:.04678-7.50 2077.85 78722797 a201 8:03: EBEN,
Width (BW) D287 274300 20775157 52341870202 3.187) 3:103:25
BL/BW 1782462241251 2072497 24525108208 25372517255
Pronotum Length (PL) Ll ve EGT 1:60-1.74 20) E89" 71:86:19 rn ZON TIEKE
Width (PW) Hy 32.0977731:99 220207 245 2402747 002.0 22802222354
PW/PL LUPE Or SEE 20 IE 127215198205 219 825419721050
PW/BW 11 0.73 0.72-0.74 20 0.77 0.76-0.78 20 0.72 0.71-0.72
Elytron Length (EL) 11 434 418-450 20 480 472-488 20 481 471-492
Width (EW) ILS ESES 0! 220) ES NS CES OMZ OM 7159915521762
EL/EW 110 0302083 0820 05 NS 003 00205 0 SUIS 06
EL/BL 11 062 061-063 20 061 0.60-0.62 20 0.60 0.60-0.60
Wing Length (WL) 1190649 CNE C2 07 SIN 5827 COR 2 00727 OTETEZG
Width (WW) MSI 923822:642 2.07 92:85 227/7528902 ON ETE 68-285
WL/WW 1192258 255222164. 201 265 2.62.2269 208 2656020
WL/EL TIRA 1451541207157. LES ATS OAN 207 151149152
Met-episternum Length (ML2) 11 0.82 0.77-0.88 20 083 081-084 20 092 0.88-0.94
Width (MW2) 11 0.61 0.58-0.63 20 0.68 0.66-0.69 20 0.65 0.63-0.66
ML2/MW?2 (ME2) MET 367 MESSE 20 AIDE ZOR 2395
Hindleg Length tarsus (TAL) 110292:00 7 7188921069207 22.092220 7A 2 0,7291 52711672775
Length tibial (TIL) 9719222108 0727187723529 252 O ZEE 2152225
TAL/TIL 11 101 099-104 20 096 094-097 20 1.01 1.00-1.02
128
AUKEMA: Three species of Calathus
Figs. 4-6. Habitus of females of Calathus. — 4, C. melanocephalus; 5, C. cinctus, 6, C. mollis. Scale 1 mm.
stede (Province of Zeeland) by the author. The
following conclusion can be drawn. Although there
are some differences between the means (e. g. in the
case of the ratio length/ width of the metepisterna),
these measurements (or ratios estimated from
them) do not allow unambiguous identification due
to significant individual variation and wide overlap
in the ranges of measurements.
Table 7 summarizes length/width ratios (MEI:
ML1/MW1 and ME2: ML2/MW2, see fig. 3) of
long and short winged males and females of cinctus
and melanocephalus, as well as of long winged
males and females of mollis (all specimens collected
or reared of the latter species turned out to be long
winged) from localities mentioned above, as well as
from a number of other localities too.
Although both ratios show a clear trend mela-
nocephalus <cinctus< mollis for both males and
females, the mutual overlap is so wide that identi-
fication on the basis of these ratios alone usually
will not be possible. The observed differences be-
tween wing-morphs and sexes are smaller than
those between species (table 7)-In general differ-
ences between species, wing-morphs and sexes are
better illustrated by ratio ME2 than by ratio MEI.
Table 8 illustrates the variation in the number of
dorsal punctures in the 3rd elytral interval of cinc-
tus and melanocephalus from Texel and mollis
from Haamstede. In both sexes and wing-morphs
of cinctus considerable deviation (>50 %) from the
standard number of three punctures in the third
elytral interval was found, whereas the deviation in
melanocephalus and mollis was relatively low (< 40
%). Both the number and position of the punctures
are also rather variable and certainly not of diagnos-
tic value.
General coloration, shape of the elytra and shape
of the right parameres are very constant and can be
used to identify most specimens according to the
key presented hereafter. Only a few long winged
females of cinctus and mollis may give some prob-
lems in identification.
129
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 7. Measurements of met-episterna: ratio length/ width. MEI: ML1/MW1; ME2: ML2/MW2 (see fig. 3); sw: short
winged; lw: long winged; N: Number of measurements.
MALES
Wing-morph Species
SW
SW
lw
FEMALES
SW
SW
lw
Locality
cinctus
melanocephalus
cinctus
melanocephalus
mollis
cinctus
melanocephalus
cinctus
melanocephalus
mollis
Nuil
Noordenveld!)
Texel
Oost-Flevoland
Nuil
Noordenveld!)
Kraloo
Bijlmermeer
Texel
Nuil
Noordenveld!)
Texel
Oost-Flevoland
Kraloo
Bijlmermeer
Texel
Meijendel
Haamstede
Nuil
Noordenveld!)
Texel
Oost-Flevoland
Nuil
Noordenveld!)
Kraloo
Bijlmermeer
Texel
Nuil
Noordenveld!)
Texel
Oost-Flevoland
Kraloo
Bijlmermeer
Texel
Meijendel
Haamstede
1): measurements M. G. J. Oude Wesselink
MEI ME2
N Range Mean 95% cl. Range Mean 95% cl.
30 106-118 1.11 110-112 1.28-1.46 136 1.35-1.38
27 10522221711 1.09-1.15 “119-47 134 131137
20 0.97-1.10 1.04 1.03-1.06 125-143 134 1.32-1.36
30 09921916, 1:0777:05-108 EO 3 ONE 29 PT
19 0.96-1.10 1.04 1.02-1.06 1.15-1.31 1.23 1.21-1.25
30 OVE AOS NO = POS NONO 25930
9) 0.98-1.10 104 1.01-1.06 1.26-1.29 1.24 1.20-1.28
gf 103-114 1.07 1.06-1.09 1.23-1.40 1.27 1.25-1.30
20 0.95-1.07 1.02 1.00-103 115-132 1.24 1.22-1.27
30 ROIO MERE EDA OO ees ele al
30 1.04-1.26 1.14 IMO AO PRINS SES JIS 1.33-1.40
11 ROO OIO DINO OIS TIEL ES Seles E139
13 1.00-1.15 1.08 105-111 114-134 124 121-128
2 1.09-1.12 1.39-1.45
3 1.07-1.09 1.21-1.29
7 1.00-1.09 1.05 1.02-1.08 1.20-1.40 1.28 1.21-1.35
14 IS 1 25 INO El EE) OA
20 alien AO) A6 MEET 1.40-1.45
20) 1.02-1.18 1.11 1.09-1.12 124-149 1.35 15352197
29 1.00-1.20 1.08 1.07-1.10 1.17-144 130 1.28-1.33
20 0.921.070 10120991025 114313027130 21272133
30 OOG OD MIDI OI ES 125 DET
30 1.00-1.10 1.04 1.03-1.05 1.14-139 1.25 1.23-1.27
30 0.98-1.22 106 104-109 1.16-139 127 1.24-1.29
12 1.00-1.13 105 103-108 118-137 1.24 1.18-1.30
10 0.98-1.18 1.09 1.05-112 1.23-1.33 1.27 1.24-1.30
20 0.92-1.07 1.01 099-102 114-130 1.22 1.20-1.24
30 KONZ 1.11-1.13 1.26-1.44 1.35 1.34-1.37
29) DOS OMAN WIG rele ASS SANA CI 31850)
11 LOB 1.08-1.14 1.25-1.49 136 131-141
30 0.97-1.13 1.06 1.04-107 1.12-141 1.26 1.24-1.28
18 0:98212187 17.087 7705-1117 1482 180551905154
7 NOMA WO, BETT LS 005112. 6216577
20 095-108 1.03 101-104 117-130 1.22 121-124
17 1.13-1.28 1.19 1.17-1.21 133-154 143 140146
20 1.05-1.24 116 115-118 129-161 142 1.39-1.45
Tabel 8. Number of punctures in 3rd elytral striae. N: number of observations; D: deviation (%) from standard number
of puntures (3/3) in third striae; sw: short winged; lw: long winged.
Punctures in 3rd elytral stria (right/left elytrum)
Species Sex Wing morph N
cinctus males sw 20
lw 11
females sw 20
lw 11
melanocephalus males Sw 20
lw 7
females sw 20
lw 20
mollis males lw 20
females lw 20
130
WO DB DL BR BIB Bie LB
fin ids bale =p Think Sue SOR?
Sin hed RT
i Rc eRe) en RR
Delbo hal sl
TU À
à e x = 5 2 E
A Rt Ind HEISA AA
on dee ee D
RZ e aan ce Pre
NE ka Md A CALL ll
4/4
DW NR be
4/5 5/3
5/4 D
= Dp)
- 545
90.0
- 63.6
= ap
- 286 |
SI)
= 300m
=, 15.00
550
Figs. 7-9. Right parameres of Calathus.—7,C. melanoce-
phalus; 8, C. cinctus; 9, C.mollis. Scale 0.5 mm.
In all three species the females are less shining
than the males, due to a denser microsculpture of
usually isodiametric meshes, which is most obvious
on the elytra.
Key to the species
1. Elytra bicoloured, with epipleura, base and dor-
sal punctures clear rufous, contrasting with the
black dorsal surface. Sides of elytra almost pa-
rallel and broadly rounded apically (fig. 4).
Pronotum more or less rufous, clearly contrast-
ing with the black head and with the elytral
surface. Sternites dark, contrasting with the ru-
fous meso- and meta-thoracical structures.
Males: right paramere widened and hooked at
the tip (fig. 7). Vesica: fig. 10. Wings reduced
or fully developed (rarely specimens with only
one fully developed wing are found) (body
IENE th (6-8 IMM) ER melanocephalus
— Elytra unicoloury rufous to piceus brown, with
sides evenly rounded (figs. 5-6). Little or no
contrast in colour between pronotum and head
and elytra, nor between sternites and meso- and
meta-thorax. Males: right paramere not wi-
dened near the apex and unarmed or with a very
small hook apically (figs. 8-9) ................. 2
2. Pronotum slightly contrasting with the darker
head and the elytra. Males: right paramere
hooked, tip unarmed or nearly so (fig. 8). Ves-
ica: fig. 11. Wings either fully developed or
strongly reduced (body length 6-8.5
ON) EEEN IE n OM Aert hd cinctus
- Pronotum not contrasting with the head and
the elytra. Males: right paramere almost
straight with a very small hook apically (fig.9).
Vesica: fig. 12. Wings always full (body length
CINTO Aaa MER a RR OM Was mollis
AUKEMA: Three species of Calathus
Habitat and life history
Habitat
Calathus melanocephalus is mainly a species of
open country, where it lives on different kinds of
moderately dry soil with sparse vegetation, achiev-
ing its greatest abundance on sandy soils. It is a
common inhabitant of dry meadows, grassland
dunes and heaths; also on agricultural land and in
thin forest, mainly of Pinus (Lindroth 1986) or
Betula (den Boer pers. comm.). Den Boer (1977) in
his subdivision of carabids in habitat groups classi-
fied melanocephalus as an F-species, i.e. mainly a
species of heath and peat moor.
Calathus cinctus is mainly found in mixed pop-
ulations with the preceding and/or the following
species in dry meadows, dune grassland, agricultu-
ral land and waste land. Van Dijk (1986) classified
cinctus as an H-species sensu den Boer (1977), i.e.
an eurytopic species, which in most cases prefer
localities significantly influenced by human activi-
ties (arable fields, pastures, farm yards, gardens and
Figs. 10-12. Vesica of Calathus. - 10, C. melanocephalus;
11, C. cinctus; 12, C. mollis. Scale 0.5 mm.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 9. Numbers and ratio of Calathus cinctus (cin) and C. melanocephalus (mel) in mixed populations.
NUIL SCHOONGELEGEN TEXEL
Species Species Species
Year cin mel %cin cin mel %cin cin mel %cin
1973 11 30 26.8 8 71 10.1
1974 13 29 31.0 3 54 5.2
1975 35 64 35.4 6 23 20.7
1976 16 20 44.4 22 39 36.1
1977 34 24 58.6 39 31 Doll
1978 59 67 46.8 172 112 60.6
1979 12 34 26.1 3 62 4.6
1980 2 19 9.5 — 40 &
1981 = 16 - - 30 =
1982 = 6 = 72 =
1983 1 = = 84 =
1984 - 6 = 50 - 82 19 81.1
1985 - 4 - 91 = 264 265 49.9
1986 - 18 - 102 = 133 310 30.0
1987 32 176 155
1988 6 74 U
Total 183 DO 253 861 Sly 844
Table 10. Weekly catches of Calathus cinctus and C. melanocephalus at Nuil, Drenthe in 1976. N: number of beetles;
ND: number of females dissected, MNE: mean number of eggs in the ovaries of dissected females.
cinctus melanocephalus
Males Females Males Females
Date N N ND Young Old MNE N N ND Young Old MNE
7 Jul 1 1 1 1 AUTO
14 Jul 5 4 4 3 1 4.3
21 Jul - 1 l 1 - 8 12 8 6 2 Wile
28 Jul = = = = = 14 18 10 9 115.9)
4 Aug - 1 1 1 2 15.0 7 10 9 8 a
11 Aug = = = = 5 11 25 ll 7 LO IS
18 Aug 1 2 2 2 - 23.0 35 26 10 7 3 22.6
25 Aug 3 2 2 1 1 35.0 46 17 10 9 1 23.0
1 Sep 13 15 15 7 8 20.2 42 34 10 8 2 25.8
8 Sep 10 16 16 9) 7 33.9 11 24 10 10 - 2165
15 Sep 21 20 20 15 5 26.5 18 21 10 10 - 1726
22 Sep 27 18 18 16 2 17.8 23 18 13 12 1 18.8
29 Sep 16 8 8 5 3 18.3 22 4 4 3 1 10.8
6 Oct 17 10 10 9 1 15.4 9 2 2 0 2 16.5
13 Oct 12 20 18 10 8 111152 6 10 10 9 Il 2.0
20 Oct 4 6 6 4 2 9.8 2 1 1 1 - -
27 Oct 2 6 6 3 3 8.0 2 4 2 2 -
3 Nov - 1 Il 1 - 1 = -
10 Nov 2 3) 1 1 - 3 1 -
Total 128 129 125 85 40 262 231 125 105 20
roadside verges). In three cases a mixed population
of cinctus and melanocephalus was observed during
a period of several years: populations at Nuil and
Dwingeloo (fields withdrawn from agricultural
practice) and at the West Frisian Island of Texel
(artificial dune grassland) (table 9). In all three
cases, however, cinctus disappeared or nearly disap-
peared after a number of years of coexistence with
melanocephalus. These data might suggest that
152
cinctus is better adapted to changing, unstable hab-
itats than melanocephalus. At Nuil and Dwingeloo
the treatments for impoverishing the amount of
nutrients (annual mowing and removal of vegeta-
tion) most likely caused the decrease of cinctus (see |
also van Dijk 1986), while on the dune grassland at |
Texel a similar treatment might be responsible for |
the same kind of shift. |
Calathus mollis is strictly confined to dry sandy
habitats in open country. It lives in sparse xerophi-
lous vegetation, common on sand dunes along the
coast, where it occurs in tufts of Elymus and Am-
mophila in company with Demetrias monostigma
Samouelle and Dromius linearis (Olivier) (Lind-
roth 1986; own observations), and only incidentally
inland, e.g. on blown sands and other large sand
deposits. In den Boer's subdivision (1977) mollis
would be classified as an E-species (species of blown
sands). In the coastal dune region as well as in
inland blown sands mixed populations of mollis
and cinctus and occasionally with both cinctus and
melanocephalus occur.
Life history
Annual rhytms. - All three species show a similar
type of development. They belong to the group of
autumn breeders with a thermic hibernation para-
pause in the larvae and a photoperiodic aestivation
parapause in the adults (Thiele 1977). As in most
autumn breeders, adults of melanocephalus are al-
most strictly night active (Thiele 1977). Larvae as
well as a varying part of the adults overwinter
(Lindroth 1986). In northern and alpine areas,
however, melanocephalus is a semivoltine spring
breeder, instead of a univoltine autumn breeder,
adapted to the shorter growing season of a cooler
climate (Forsskähl 1972, DeZordo 1979, Refseth
1988). Under these circumstances larvae of mela-
nocephalus need almost two years to develop.
Reproduction period and age-composition. - De-
tailed information about the life history of mela-
nocephalus is given by Vlijm & van Dijk (1967),
Vlijm et al. (1968) and van Dijk (1972, 1973).
Adults emerge in June and July, reproduce from
August onwards and a varying part of these beetles
hibernate to become active again in May-June, re-
producing for the second time from July onwards.
The total reproduction period covers the months
July, August and September. Vlijm & van Dijk
(1967) and Vlijm et al. (1968) estimated the second
year category as about one third of the adult pop-
ulation at Schiermonnikoog, but this figure seem to
show significant annual and local variation (van
Dijk 1972, Baars & van Dijk 1984). Even three and
four year old beetles were found by a mark-recap-
AUKEMA: Three species of Calathus
mean number of eggs laid
o cinctus
O melanocephalus
A mollis
30 35 40 45 50
August September October November December
week number/month
Fig. 13. Mean egg production of Calathus cinctus (29
females), C. melanocephalus (23 females) and C. mollis
(16 females) and mean weekly temperature in 1981 under
outside conditions.
ture experiment in a populations of melanocepha-
lus (van Dijk 1979b), representing 26.5 and 0.5 %
of the total catch during four years respectively.
Calathus cinctus was found to reproduce in
Drenthe somewhat later than melanocephalus (Au-
gust-November) (van Dijk 1978). From weekly pit-
fall samples of cinctus collected in 1976 at Nuil, 165
females caught between 21 July and 17 November
were dissected and checked for their ovarian devel-
opment. The main reproductive period, defined
Table 11. Reproduction period of Calathus cinctus, C. melanocephalus and C. mollis in outdoor breeding experiments.
N: number of females.
REPRODUCTION PERIOD
Year Species N Egg-laying period Mean number of eggs> 10
1981 cinctus 29 12 Aug - 9 Dec (18 weeks) 26 Aug - 21 Oct (9 weeks)
melanocephalus 36 31 Jul —6 Nov (15 weeks) 7 Aug - 9 Oct (10 weeks)
mollis 14 3 Aug - 7 Dec (19 weeks) 31 Aug - 19 Oct (8 weeks)
1982 cinctus 30 16 Aug - 31 Jan (25 weeks) 30 Aug - 22 Nov (13 weeks)
melanocephalus 8 30 Jul - 22 Oct (13 weeks) 13 Aug - 15 Oct (10 weeks)
mollis 17 6 Aug- 3 Jan (23 weeks) 20 Aug- 5 Nov (12 weeks)
133
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
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mean number of eggs laid
30 35 40 45 50 1
August September
October November December January
AUKEMA: Three species of Calathus
Fig. 14. Mean egg production of Cala-
thus cinctus (30 females), C. melanoce-
CI - 1993 ne DIA rome) and < mollis au
females) and mean weekly tempera-
tures in 1982/1983 under outside con-
ditions.
o cinctus
O melanocephalus
4 mollis
February
week number /month
here as the period in which the mean number of
eggs in the ovaries is 10 or more, fell between the
first week of August and the third week of October
(table 10). Forty females (24.2 %) were classified
as old females, reproducing for the second year.
Comparable data for melanocephalus collected at
the same site resulted in a main reproductive period
between the third week of July and the first week
of October and 16 % old females (table 10), of
which only a single female caught on 13 October
was not or no longer reproductive.
Calathus mollis was not studied in detail in this
respect but hand catches on the West Frisian Island
of Ameland on 27 May 1981 comprised old beetles
only (29), whereas on 28 June 1979 about 12 % of
the 248 beetles collected at the same site were just
emerged tenerals. These data suggest a similar type
of age composition of populations of this species.
In the laboratory a small part of the females of
all three species kept individually even survived a
second hibernation period and laid eggs during a
third year.
Outdoor breeding experiments carried out in
1981 and 1982 also gave an impression of the re-
productive period of the three species (table 11).
Also in these experiments melanocephalus showed
an earlier start and finish of the egg-laying period
than both cnctus and mollis. Differences between
the results for the two years are attributed to differ-
ent temperature conditions. In 1981, for instance,
all females stopped egg production in the first third
of December because of a sudden frost period (fig.
13). Due to the later start, cinctus and mollis suf-
fered more from this event than melanocephalus.
Under the more favourable weather conditions of
1982/1983 melanocephalus showed about the same
reproduction period, whereas both cinctzs and mol-
lis were able to prolong their reproduction periods
substantially under these conditions (fig. 14).
Data on egg production and larval development
of the three species in relation to wing develop-
ment, temperature and availability of food will be
published elsewhere.
Daily rhythms. — Information about the daily
activity of the species was derived from a small
experiment in which the egg production was esti-
mated every two hours under fixed temperature and
daylength conditions. On 14 September 1981 eight
reproductive females of cinctus, melanocephalus
and mollis, collected at Ameland on 14 July, were
transferred from the outdoor breeding house to an
incubator adjusted at 19°C and a daylength of 14.75
hours (6.15-19.00 h). They were kept there with
135
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
males in single pairs for three days. In all three
species egg production took place mainly during the
dark hours (18.30-6.30 h); a very few eggs were also
laid during the early morning hours (table 12).
Emergence of adults. - The outdoor breeding
experiments also gave an impression of the emer-
gence period of the adults (table 13). In 1982 almost
all cinctus, melanocephalus and mollis emerged in
the first two thirds of June, and in 1983 in the latter
half of the same month. Van Dijk (1973) trapped
teneral females of melanocephalus in the field as
well as in enclosure experiments during a short
period from 15 June onwards. Samples of cinctus
and mollis collected on 28 June 1979 on the West
Frisian Island of Ameland consisted of 75 % (102
of 138 specimens) and 11.7 % (26 of 248 speci-
mens) tenerals respectively.
Feeding. - Feeding behaviour and prey spectrum
of melanocephalus were studied by Hengeveld
(1980) by identifying food remains in the gut con-
tents of beetles trapped in the field and killed in-
stantly. From the diet components found, it ap-
peared that melanocephalus is a polyphagous, but
not exclusively carnivorous species (see also Melber
1983). Similar results were found by Smit (1957),
Skuhravy (1959) and Hengeveld (1980) for the
closely related Calathus ambiguus (Paykull), C. er-
ratus Sahlberg and C. fuscipes (Goeze).
Dispersal. - Den Boer (1977) developed criteria
to classify carabid beetles in three dispersal groups
using data on wing development, wing-morph fre-
quencies and actual flight (catches of flying beetles
in window traps). A-species are those with a rela-
tively low power of dispersal (monomorph bra-
chypterous or wing-polymorphic species with a
very low frequency (< 1%) of macropterous bee-
tles). B-species have a relatively high power of
dispersal due to their capability of flight (macro-
pterous and wing-polymorphic species caught in
window traps, see also van Huizen 1980), whereas
C-species are the remaining species (macropterous
Fig. 15. Distribution of Calathus cinctus in The Nether-
lands. Symbols on this and the following maps: open
circles: recorded before 1930 only; small dots: recorded
between 1930 and 1960; large dots: recorded since 1960.
or wing-polymorphic with a high frequency of ma-
cropterous beetles, but never or rarely caught fly-
ing) with uncertain dispersal power.
The wing-dimorphic cinctus was caught in win-
dow traps (van Huizen 1980; den Boer et al. 1980)
and consequently belongs to the group of B-species
(van Dijk 1986).
The wing-dimorphic melanocephalus and the
monomorph macropterous mollis were never
caught flying, but laboratory-bred beetles of both
species developed flight muscles and flew fre-
quently in a laboratory test (to be published else-
where). For the time being both species therefore
have to be considered C-species. Den Boer (1977)
Table 13. Emergence of adult beetles in outdoor breeding experiments. N: number of beetles.
SPECIES
cinctus melanocephalus mollis
Year Decade N N N
1982 1-10 June 3 ( 1.8 %) 113 (43.3 %) 158 (98.7 %)
11-20 June 136 (80.5 %) 101 (38.7 %) 2 (I Gy)
21-30 June 29 (17.2 %) 40 (15.3 %) = =
1-10 July l ( 0.6 %) 7 (27%) = =
1983 1-10 June 2 > = 3 a =
11-20 June 69 (53.9 %) 12 (34.3 %) 32 (68.1 %)
21-30 June 58 (45.3 %) 23 (65.7 %) 15 (31.9 %)
1-10 July 1 ( 0.8 %) = = = =
136
Fig. 16. Distribution of Calathus melanocephalus in The
Netherlands.
and van Dijk (1986) treated melanocephalus as A-
species, based on the extremely low frequency of
macropterous beetles in populations investigated in
the Dutch province of Drenthe (0.23 %). Popula-
tons in the recently reclaimed Flevopolders, how-
ever, showed a considerable higher freqency of ma-
cropterous beetles (25-40%: den Boer 1970; Auke-
ma 1986).
During breeding the flight muscles are autolysed
in these species and flight only occurs before breed-
ing. Data on the development of flight muscles and
the occurrence of actual flight from the field and
from breeding experiments in relation to temper-
ature and availability of food will be treated else-
where (Den Boer et al. (1980) already gave some
preliminary data for cinctus).
Distribution
Distribution in The Netherlands
Available data were used to prepare distribution
maps on the UTM 10 X 10 square km scale as used
by the European Invertebrate Survey in The Ne-
therlands (figs. 15-17).
Calathus cinctus (fig. 15) and C. melanocephalus
(fig. 16) are widely distributed over the whole coun-
try, whereas C. mollis (fig. 17) is almost restricted
to the coastal dune region, with only a few inland
populations on blown sands or other large sand
deposits. Desender (1985, 1986) and Moncel (1972)
AUKEMA: Three species of Calathus
Fig. 17. Distribution of Calathus mollis in The Nether-
lands.
published comparable maps and data for Belgium
and France respectively.
A closer examination of the data of cinctus (fig.
15) shows, that this species has been caught regu-
larly in The Netherlands since 1864 (Overveen, vi,
1 female, leg. J. Kinker, coll. Zodlogisch Museum,
Amsterdam). For melanocephalus and mollis re-
spectively the first records are Overveen, x.1864, J.
Kinker, 1 male (coll. Zodlogisch Museum, Amster-
dam) and Driehuizen, 23.vii.1882, 1 male (coll.
Zoölogisch Museum, Amsterdam).
Time-trends in the number of 10 X 10 km
squares occupied per decade by the three species
were estimated according to the method of Turin &
den Boer (1988). Fig. 18 shows the time-trends
after correction of the number of occupied squares
for differences in collecting intensity between the
decades (table 14). Calathus melanocephalus shows
a significantly increasing time-trend, whereas both
cinctus and mollis show rather indifferent time
trends (fig. 18). When considering the 20th century
only (table 14: decades 4-12) all three species show
negative correlation coefficients, but only in the
case of cinctus was this significant, largely due to the
high number of squares occupied in the years 1910-
1930.
Ranges
Concerning their general distribution, revision of
material on a larger scale is necessary to get a
15%
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 14. Number of 10 x 10 km squares occupied per decade by Calathus cinctus, C. melanocephalus and C. mollis. TO:
total number of squares occupied by all carabid species per decade; F: correction factor calculated from the total number
of squares covering The Netherlands (F=448/TO); O: actual number of occupied squares; OF: corrected number of
occupied squares (OF=O%F).
Decade cinctus melanocephalus mollis
Nr Years TO F O OF O OF O OF
1 1870 - 1879 72 6.22 2 12 0 0 2 12
2 1880 - 1889 81 399) 1 6 1 6 3 17
3 1890 - 1899 108 4.15 5 21 7 29 5 21
4 1900 - 1909 150 3.03 16 48 18 55 6 18
5 LOOS IOI 1972 255 26 61 24 56 11 26
6 1920-1929 214 2.10 22 46 28 59 9 19
7 1950219 95215 2.10 18 38 29 61 15 32
8 1940 - 1949 246 1.82 12 22 26 47 7 15
9) 1950-1959 264 1.70 16 27 35 59 5 22
10 1960 - 1969 312 1.44 9 13 20 29) 10 14
Ul 1970-1979 336 135 23 31 35 47 12 16
12 1980 - 1989 350!) 1.28 21 27 33 42 9) 11
') Estimated value. Up till now 14714 records from 321 squares were recorded, but many have to be added (H. Turin,
personal communication)
reliable picture (the data in Turin (1981) are only
based on literature). In the Dutch collections men-
tioned and/or in the collections of the British Mu-
seum (Natural History), London, the Zoologisches
Museum, Berlin, the Zoologische Staatssammlung,
Miinchen, and M. Baehr, Miinchen, I have seen
material from the following localities:
C. cinctus: South-England (south of the line
River Seven -The Wash), Wales, Ireland, The Ne-
therlands, Belgium, Germany, Poland, the Baltic
States (Estonia), Russia, France, Spain, Sardinia,
Corsica, Italy, Sicily, Malta, Yugoslavia, Austria,
Hungary, Romania, Bulgaria, Albania, Greece, Cy-
prus, Turkey, Crimea, Ukraine, the Caucasus, Iran,
Syria, Israel and Morocco;
C. melanocephalus: Finland, Sweden, Norway,
Denmark, Iceland, England, Scotland, Wales, Ire-
land, The Netherlands, Belgium, Luxembourg, Ger-
many, Poland, the Baltic States (Estonia), France,
Spain, Sardinia, Corsica, Italy, Sicily, Yugoslavia,
Switzerland, Austria, Hungary, Romania, Albania,
Greece, Turkey, Ukraine and the Caucasus;
C. mollis: Denmark, England, Wales, Scotland,
Ireland, The Netherlands, Belgium, Germany (the
East Frisian Islands), France, Portugal, Spain, Mal-
lorca, Italy, Sicily, Yugoslavia, Greece, Turkey, Gi-
braltar, Malta, Algeria and Morocco.
Lindroth (1986) indicated the occurrence of cırc-
tus in South-Sweden and Denmark (see also Bangs-
holt 1983) and of mollis in Norway. Since Lindroth
listed the right combination of characters for both
taxa, his findings, although not confirmed, are con-
sidered reliable. Anderson (1987, in litt. 1990)
furthermore found cinctzs to be sporadically dis-
tributed around the Irish coast south of the line
from the Shannon Estuary to Dublin.
Calathus melanocephalus obviously has a Euro-
138
number of corrected squares
60 cinctus
40
20
melanocephalus
60
40
20
mollis
60
40
20
oO
N
-
(CA
oo
2
decade
Fig. 18. Plots of the corrected time trends of Calathus
cinctus, C, melanocephalus and C. mollis; decades: 1870-
1879 (1), 1880-1889 (2), 1890-1899 (3), etc.
pean distribution, ranging from Iceland and Scan-
dinavia in the North to the Mediterranean in the
South. It is the only species of this group also oc-
curring at higher altitudes in mountain areas. The
subspecies alpinus Dejean is restricted to the east-
ern Alps (Koch 1989). Ball & Negre (1972) list a
probably introduced specimen of melanocephalus
from North America.
Calathus cinctus does not occur as far north as
melanocephalus, but in the Mediterranean it
reaches further south, occurring as far as Morocco
and Algeria in the west and Syria and Iran in the
east.
Calathus mollis seemingly is restricted to the
Atlantic and Mediterranean coastal areas.
DISCUSSION
The establishment of Calathus (Neocalathus)
cinctus as a good species has solved a major iden-
tification problem in the genus Calathus, although
even now long winged females of cinctus are diffi-
cult to distinguish from those of mollis. On the
other hand, a new problem arises in the interpre-
tation of existing literature on the species of the
melanocephalus-group. It is quite clear from the
data presented here that cnctus, although in liter-
ature still considered to be mainly distributed in
Eastern Europe (southeast and east of the river
Weser) (Lienemann 1981; Koch 1989), is a very
common species in Western Europe too. For in-
stance in England circtus was already present at the
time of Stephens, because in his collection, bought
in 1853 by the British Museum (Natural History),
London, and kept in its original state (Hammond
1972) four out of the 13 specimens in the box of
melanocephalus belong to cinctus (1 male and 3
females, all short winged). Furthermore Jeannel
(1942) already indicated the occurrence of cinctus in
western France (Vendée).
Lindroth (1949) used data on wing development
in Calathus mollis as one of the main illustrations
for his theory of postglacial recolonization of Fen-
noscandia. Although Lindroth indicated in the same
publication that there were two taxa involved, his
data were never revised. Also the data from the
following studies will have to be reconsidered con-
cerning species identity: for larval taxonomy
Table 15. Spearman rank correlation coefficients be-
tween the corrected number of squares occupied and the
decade rank
Species All decades 20th Century!)
cinctus 021 -0.74*
melanocephalus +0.72* -0.54
mollis -0.20 -0.60
*) 0.02 <P<0.05
!) decades 4-12 of table 14
AUKEMA: Three species of Calathus
(Kurka 1971: description of larval stages of mela-
nocephalus and mollis), for diagnostics of eggs
(Luff 1981: characteristics of the eggs of melanoce-
phalus and mollis), for chromosome numbers (Ser-
rano 1981: chromosome numbers of melanocepha-
lus (n=37) and mollis (n=39)), for population
dynamics (Gilbert 1956: population dynamics of
melanocephalus and mollis in the coastal dunes of
the Isle of Anglesey where, as we know now, all
three species occur), for the occurrence in agricul-
tural fields (Luff 1987: only references for the oc-
currence of melanocephalus), for flight observa-
tions (Palmén 1944: long-winged melanocephalus
washed ashore on the southwestern coast of Fin-
land; Honek & Pulpán 1983: melanocephalus
caught ina light trap), and for bionomics and breed-
ing (Kurka 1972: melanocephalus and mollis).
CONCLUSIONS
1. Calathus (Neocalathus) cinctus Gemminger &
Harold is considered a good species, reproduc-
tively isolated from both the closely related C.
(N.) melanocephalus (Linnaeus) and C. (N.)
mollis (Marsham).
2. The species of this so-called melanocephalus
group can be distinguished from each other
reliably only on the basis of the shape of the
elytra, coloration characters and the shape of
the right parameres.
3. Morphological measurements and ratios are
not useful in identifying these species.
4. Both cinctus and melanocephalus are wing-di-
morphic with either fully developed wings (of
about 1.5 times the length of the elytra) or
reduced wings (of about 0.5 times the length of
the elytra), whereas Dutch mollis always are
macropterous.
5. According to the subdivision in habitat groups
of Den Boer (1977) cinctus is classified as H-
species (species preferring localities signifi-
cantly influenced by human activities), mela-
nocephalus as mainly a species of heath and
peatmoor (F-species) and mollis as E-species
(species of blown sands/dunes).
6. The species are night-active autumn breeders.
Calathus melanocephalus reproduces earlier
and during a shorter period in the season than
cinctus and mollis.
7. Age-composition of populations of cinctus and
mollis resembles those of melanocephalus (not
only larvae, but also a varying part of the adult
beetles overwinters and reproduces for a second
time).
8. Concerning dispersal power cinctus is classified
as a good disperser (B-species sensu Den Boer,
1977), whereas melanocephalus and mollis are
considered C-species (species with uncertain
159
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
dispersal power) by lack of information on ac-
tual flight.
9. All three species are common in The Nether-
lands and were so during the 20th century. Both
cinctus and melanocephalus are well distributed
all over the country, whereas mollis is limited
to coastal dunes and a few inland blown sands
or other large sand deposits.
10. All three species are mainly european in distri-
bution, with melanocephalus reaching further
north and occurring also at higher altitudes in
mountain areas, cinctus reaching further south
and mollis limited to coastal areas in the Atlan-
tic and mediterranean area.
ACKNOWLEDGEMENTS
Many thanks are due to P. J. den Boer, J. C. van
Lenteren and M. L. Luff for critical comments on
the manuscript, to Th. S. van Dijk, Y. Jongema, A.
Spee, T. H. P. van Huizen, M. G. J. Oude Wesselink,
J. Popping and H. Lindenschot for their assistance
during practical work, to P. Hammond and M. J. D.
Brendell (British Museum (Natural History), Lon-
don), M. Baehr (Zoologische Staatssammlung,
München), F. Hieke (Zoologisches Museum, Ber-
lin), J. Krikken (Rijksmuseum van Natuurlijke His-
torie, Leiden), B. Brugge and W. Hogenes (Zoölo-
gisch Museum, Amsterdam) and J. G. Rozeboom
(Department Entomology, Agricultural University,
Wageningen) for the loan of specimens and to all
those mentioned under material for supplying ad-
ditional material. M. L. Luff improved the English
text.
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nikoog. — Zeitschrift für Morphologie und Okologie
der Tiere 58: 396-404.
Vlijm, L., Th. S. van Dijk & S. Y. Wijmans, 1968. Ecolog-
ical studies on carabid beetles. III. Winter mortality in
adult Calathus melanocephalus (Linn.). Egg produc-
tion and locomotory activity of the populations which
has hibernated. - Oecologia 1: 304-314.
Wiebes-Rijks, A. A., 1959. Het onderscheid tussen Cala-
thus melanocephalus L. en Calathus mollis Marsh. (Co-
leoptera, Carabidae). — Entomologische Berichten,
Amsterdam 19: 248-253.
Received: 13 February 1990
Accepted: 16 August 1990
141
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
142
J. BELLE & J. van TOL
National Museum of Natural History, Leiden
ANOMALAGRION HASTATUM (SAY), AN
AMERICAN DAMSELFLY INDIGENOUS TO THE
AZORES (ODONATA, COENAGRIONIDAE)
Belle, J. & J. van Tol, 1990. Anomalagrion hastatum (Say), an American damselfly indigen-
ous to the Azores (Odonata, Coenagrionidae). - Tijdschrift voor Entomologie 133: 143-147,
figs. 1-16. [ISSN 0040-7496]. Published 14 December 1990.
Anomalagrion hastatum (Say), a species hitherto known only from the New World, is
recorded from the European archipelago of the Azores. It was captured for the first time
on these islands in 1938, but was mistaken for Ischnura senegalensis (Rambur). Only
females have been found up to now, and it is hypothesized, that this species reproduces
parthenogenetically.
Correspondence: Dr. J. Belle, Onder de Beumkes 35, 6883 HC Velp, The Netherlands.
Key words. - Azores; Anomalagrion; Ischnura; Europe; parthenogenetic reproduction.
In 1940 Valle published a paper on the Odonata
of the Azores and Madeira, in which the occurrence
of Ischnura senegalensis (Rambur) was reported
for both the Azores and the Canary Islands. Belle
(1982) examined the specimens in question from
the Azores. He found that they were characterized
by a very small vulvar spine and he expressed his
doubts on the correctness of the identification, since
the vulvar spine of I. senegalensis characteristically
is very long (figs. 12, 15). His observations on the
Canary Islands only revealed Ischnura sabarensis
Aguesse, 1958. Hämäläinen (1986) could confirm
Belle's supposition that Valle’s identification for the
individuals from the Canary Islands was incorrect.
All specimens belonged, indeed, to Ischnura saha-
rensis.
The identity of the specimens of the Azores,
however, remained uncertain due to the fact that
only females were available. In the summer of 1988,
the senior author decided to visit the Azores him-
self in order to trace the male of the misdetermined
Ischnura species. He visited the Isles of Sào Miguel,
Sao Jorge and Pico. On Sao Miguel and Sao Jorge,
the desired species was not found, but on Pico more
than 30 females were captured. Despite diligent
hunting, the matching male was not encountered.
To unravel the status of the collected females,
they then were handed over to the junior author for
examination. After consultation of literature and
the collection of the National Museum of Natural
History, Leiden (RNHL), it soon became obvious,
that they did not belong to one of the Ischnura
species known from Europe or North Africa. They
were referable to the closely related New World
genus Anomalagrion, and identified as Anomal-
agrion hastatum (Say). This identification was also
shared by Drs. R. W. Garrison (Azusa, USA) and T.
W. Donnelly (Binghamton, USA), who in collabo-
ration examined three of the females.
Anomalagrion hastatum (Say, 1839)
(Figs. 1-6, 11)
Selected references
Agrion hastata Say, 1839: 38-39.
Ischnura (Anomalagrion) hastatum. - Selys 1876: sep. 11-
15?
Anomalagrion hastatum. — Calvert 1901-1908: 130-131,
390; Garman 1927: 39-40, figs. 9-10, plate 2 (7);
Walker 1953: 275-278, plate 11 (4), 21 (4), 26 (10), 30
(76
Ischnura senegalensis Rambur sensu Valle 1940: 2, 4
[misidentification].
Material
Azores. Pico: Lagoa do Caiado, 23 July 1988, 13 females;
26 July 1988, 12 females; Lagoa do Capitào, 24 July 1988,
7 females (J. Belle) in RNHL, but 3 specimens in coll.
Garrison and one specimen in the collection of the Uni-
versity of Ghent. — Other material examined (all in
RNHL): USA (Indiana, Texas, Georgia), Suriname, Gala-
pagos Islands. Also the specimens mentioned by Valle
(1940) have been examined.
Description
As indicated above, only females were collected.
The material is remarkable homogeneous for a var-
iable species as Anomalagrion hastatum appears to
be in America. Although the specimens are rela-
tively large for this species, they are very small for
143
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
144
an European damselfly. They fall within the range
of the smallest European species, Nehalennia spe-
ciosa (Charpentier). All specimens seem to be ma-
ture.
Head (fig. 1). - Labium creamish white, labrum
light brown, medio-basally with a brownish black,
triangular depression (not present in specimens
from the New World, available in the RNHL col-
lection); anteclypeus creamish white, postclypeus
shiny black; frons and genae yellowish white; dor-
sum of head dull black, the posterior spots not or
only just discernable, but hind margin of occiput
bordered with a pale stripe, connecting the pale
areas of the rear side of the head. Head covered with
fragile, pale setae.
Thorax. - Prothorax (fig. 2) predominantly dull
black, but sides yellowish white; hind margin
simply built with a rectangular median projection.
Synthorax with dorsal side black, but otherwise
light brown and creamish. Humeral suture indi-
cated in black. Mesostigmal laminae simply built
(fig. 3) with a posterior ridge, rounded on lateral
side. Lateral one-fourth of mesostigmal laminae
pale coloured. Legs rather short, creamish white,
but outerside of all femora black, and basal two-
thirds of fore tibiae with a black stripe. Wings
rather short, in rest approximately reaching the
hind margin of segment 6; latero-posterior side of
pterostigma (fig. 4) rounded with the two cross-
veins behind it very close to each other.
Abdomen. - Dorsum of all segments rather dull
black with some bluish of greenish shine and not
interrupted on any segment, nor any trace of blue
coloration, as e. g. in Ischnura sabarensis. Posterior
part of segment 10 not conspicuously raised. Vulvar
spine very small, hind margin of sternite 8 not
elongated or otherwise produced posteriad (figs. 5-
6).
COMPARISON WITH EUROPEAN SPECIES OF
ISCHNURA
At least three palaearctic species of the genus
Ischnura have to be considered for the archipelago,
viz. I. pumilio (Charpentier), I. senegalensis and
Ischnura saharensis Aguesse. The first species is
widespread in Europe, and not uncommon on the
Azores (Valle 1940, observations by J. Belle), the
second one was recorded from the Azores (Valle
1940) and from the Canary Islands (Valle 1955).
However, all specimens of Ischnura senegalensis
Figs. 1-10. — 1, Anomalagrion hastatum (Say), female
from Pico, head, dorsal; 2, idem, prothorax, dorsal; 3,
idem, laminae mesostigmalis; 4, idem, pterostigma; 5-6,
idem, end of abdomen, left profile; 7, Ischnura senegalen-
sis (Rambur), female from W. Java, laminae mesostigma-
lis; 8, Hind margin of prothorax of female, Ischnura genet
(Rambur) from Sicilia; 9, idem, I. pxmulio (Charpentier)
from Pico; 10, idem, I. saharensis Aguesse from Algeria.
BELLE & VAN TOL: Anomalagrion
recorded from the Canary Islands appeared to be-
long to I. saharensis (Belle 1982, Hämäläinen
1986). We also found that specimens from the
Azores identified as I. senegalensis by Valle evi-
dently belonged to another taxon. Apart from these
species already recorded from the atlantic islands,
also the other species of Ischnura known from
northern Africa and southern Europe, have to be
taken into account, viz. I. graellsii (Rambur), a com-
mon species on the Iberian peninsula, I. fountainei
Morton, a species known from northern Africa and
the Middle East, and less likely also I. elegans
(Vander Linden), the most common European spe-
cies and I. genei (Rambur), known from Corsica,
Sardinia, Sicilia and Malta.
Reliable characters for distinguishing females of
Ischnura and related genera are found in the shape
of the hind margin of the prothorax (figs. 8-10), the
mesostigmal laminae (fig. 7), and the vulvar spine
(figs. 11-16). In many species identification is ham-
pered by the presence of two female forms, one
similar to the male (homeomorphic) and one quite
dissimilar (heteromorphic) form. This pheno-
menon was extensively described by Schmidt
(1967).
The hind margin of the prothorax of Ischnura
species is provided with two transversal ridges,
which are developed in various ways in the Euro-
pean Ischnura species. Ischnura senegalensis males
(and the homeomorphic females) have two sub-
equal, parallel, unconnected ridges. Although this
character is unique to /. senegalensis, it is somewhat
confusing that the heteromorphic females are more
like the species of the I. elegans group. The females
pertaining to the I. elegans group (with I. elegans,
I. genet, I graellsu and I. sabarensis) can be recog-
nized by the conformation of the raised hind mar-
gin of the prothorax having a longer or shorter
anterior ridge developed into a protuberance. The
raised part of the posterior ridge is most luxuriantly
developed in I. genez, which also has a more or less
incised protuberance on the anterior ridge (fig. 8).
The protuberance is longer and the posterior ridge
shorter in I. elegans, so that the protuberance of the
anterior ridge almost fully covers the upraised part
of the posterior ridge. The hind margin of the
protuberance is usually more or less squarish (e.g.
Geijskes & Van Tol 1983, fig. 121, Askew 1988, fig.
138), but female specimens from southern France
tend to have a more incised hind margin of the
protuberance. Although the ground plan of the pro-
thorax of I. saharensis is similar to that of I. elegans,
the protuberance of the anterior ridge is hardly
developed and the whole appearance is determined
by the upraised hind margin of the posterior ridge
(cf. Aguesse 1958, fig. 3).
The hind margin of the prothorax of females of
Anomalagrion is quite dissimilar, since it is simply
built without subparallel ridges. The males of A.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 11-16. — Abdomen of females in ventral (figs. 11-14) and left lateral (figs 15-16) view. - 11, Anomauagrıon
hastatum (Say) from Pico; 12, 15, Ischnura senegalensis from W. Java; 13, Ischnura pumilio (Charpentier) from the
Azores; 14, Ischnura graellsii (Rambur) from Spain, Cadiz; 16, Ischnura saharensis Aguesse from Gran Canaria.
hastatum are absolutely unmistakable, since they
are characterized by a pterostigma in the forewing
that is retracted from the anterior margin of the
wing. They also have a greatly enlarged cylindrical
bifid process on the tergum of segment 10, but that
character occurs in a less conspicuous form also in
some North-American Ischnura species. The fe-
males of A. bastatum also differ in several charac-
ters from Ischnura species. The differential features
were already described by de Selys (1876)... ‘The
females are more difficult to identify, since they
have a normal pterostigma. They are, however,
146
quite easily distinguishable by the coloration of the
head and the thorax, which are both similar to that
in the male, by the spine of segment 8, which is
much shorter than those in Ischnura, and, finally,
on segments 8 and 9, which are black (which is
exceptional in Ischnura).... He also mentions the
existence of a orange coloured variety, which is the
immature stage (Calvert 1907: 390).
Recently, a second species in the (sub)genus
Anomalagrion was described (De Marmels 1987).
Although the pterostigma of the male forewing of
this species is greatly enlarged, it is not retracted
from the anterior margin of the wing. Attribution
to the (sub)genus Anomalagrion should therefore
be considered preliminary, awaiting a revision and
phylogenetic analysis of Ischnura and related
genera. Given the autapomorphies of Anomal-
agrion hastatum, a more than superficial analysis is
needed to settle its position in the Coenagrionidae.
BIOLOGICAL OBSERVATIONS
It is most remarkable that no males were col-
lected on the Azores up to now. The senior author
spent in three days collecting many hours in the
field without any result. We now hypothesize, that
Anomalagrion is, at least on the Azores, a parthe-
nogenetically reproducing species.
Several other characters support this hypothesis.
Firstly, the morphology of the specimens of the
Azores is very equal, while the specimens from
localities wide apart in North and South America
are quite dissimilar. This phenomemon can be un-
derstood for species with (partly) parthenogenetic
reproduction, where there is no or hardly any gene-
flow between populations, and specimens on one
locality are the offspring of only one female. Se-
condly, it seems that in North America Anomal-
agrion hastatum is a species frequenting temporary
or recently established habitats with females much
more common than males. It is obvious that fully
or partially parthenogenetic reproduction is advan-
tageous to species of this kind of habitat, while also
the likelihood to produce offspring after incidenti-
cal transport by air to a remote island is signifi-
cantly enlarged. Further biological studies are
needed to test our hypothesis. As far as we know,
there is no earlier record of parthenogenetic repro-
duction in Odonata.
The occurrence of an American dragonfly species
on the Azores is consistent with the composition of
the avifauna of these islands. There is a remarkably
high percentage of American bird species among
the migrant visitors. Den Hartog & Lavaleye
(1981), for instance, could recognize four American
species among the 23 species of non-breeding birds
seen during a six weeks stay on six islands of the
Azores (cf. Bannerman & Bannerman 1966).
ACKNOWLEDGEMENTS
We gratefully acknowledge the expert help of
Drs. R. W. Garrison (Azusa, USA) and T. W. Don-
nelly (Binghamton, USA) to check our identifica-
tion of the Anomalagrion species of the Azores. Mr.
J. C. den Hartog (National Museum of Natural
History, Leiden) gave valuable advise on the avifau-
nal composition of the Azores.
BELLE & VAN Tor: Anomalagrion
REFERENCES
Aguesse, P., 1958. Une sous-espèce nouvelle d’Ischnura en
Afrique du Nord. - Revue française d’Entomologie 25:
149-157, figs. 1-5, map 1, tabs. 1-2.
Askew, R. R., 1988. The dragonflies of Europe. - Harley
Books, Colchester: 1-291, figs. 1-502, plates 1-29.
Bannerman, D. A. & W. M. Bannerman, 1966. Birds of the
Atlantic Islands. Volume 3. A history of the birds of the
Azores. - Oliver & Boyd, London: i-xx + 1-262.
Belle, J., 1982. Odonata collected in the Canary Islands. —
Entomologische Berichten, Amsterdam 42: 75-77.
Calvert, P. P., 1901-1908. Insecta. Neuroptera. Odonata. —
Biologia Centrali Americana: i-xxx + 17-420 + plates
2-10.
De Marmels, J., 1987. Ischnura (Anomalagrion) cruzi sp.
n., eine neue Kleinlibelle aus Kolumbien (Odonata:
Coenagrionidae). - Mitteilungen der Entomologischen
Gesellschaft Basel (N. F.) 37: 1-6, figs. 1-13.
Garman, P., 1927. Guide to the insects of Connecticut. Part
V. The Odonata or dragonflies of Connecticut. — Bul-
letin of the State Geological and Natural History Sur-
vey, State of Connecticut 39: 1-331, figs. 1-67, plates 1-
22 excl.
Geijskes, D. C. & J. van Tol, 1983. De libellen van Neder-
land (Odonata). - Koninklijke Nederlandse Natuur-
historische Vereniging, Hoogwoud: 1-368, figs. 1-539.
Hartog, J. C. den & M. S. S. Lavaleye, 1981. Birds obser-
vations in the Azores (12 September-1 November
1979). - Bocagiana 56: 1-19 + plates 1-8, tabs. 1-2.
Hämäläinen, M., 1986. Note on misidentification of the
first Zygoptera material from the Canary Islands. —
Notulae Odonatologicae 2: 131-132.
Peters, G., 1988. Libellen (Odonaten) von den Kanari-
schen Inseln. - Entomologische Nachrichten und Be-
richte 32: 39-40.
Say, T., 1839. Descriptions of new North American neur-
opterous insects, and observations on some already
described. — Journal of the Academy of Natural Scien-
ces of Philadelphia 8: 9-46.
Schmidt, E., 1967. Versuch einer Analyse der Ischnura
elegans-Gruppe (Odonata, Zygoptera). - Entomolo-
gisk Tidskrift 88: 188-225, figs. 1-2, tabs. 1-5, plates 1-
7,1 map.
Selys Longchamps, E. de, 1876. Synopsis des Agrionines,
Sme légion: Agrion (suite). Le genre Agrion. - Bulletin
de l'Académie royale de Belgique (series 2) 41/42: sep.
1-282.
Valle, K. J., 1940. Odonaten von den Azoren und Madeira.
— Commentationes Biologicae 8 (5): 1-7.
Valle, K. J., 1955. Zygopteren (Odonata) von den Kanar-
ischen Inseln. - Annales Zoologici Fennici 21: 182.
Walker, E. M., 1953. The Odonata of Canada and Alaska.
Volume 1. - University of Toronto Press, Toronto: i-
xi + 1-292, plates 1-44.
Received: 28 September 1990
Accepted: 20 October 1990
147
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
148
P. K. CHAUDHURI & S. CHATTOPADHYAY
Department of Zoology, University of Burdwan
CHIRONOMIDS OF THE RICE PADDY AREAS OF
WEST BENGAL, INDIA (DIPTERA: CHIRONOMIDAE)
Chaudhuri, P. K. & S. Chattopadhyay, 1990. Chironomids of the rice paddy areas of West
Bengal, India (Diptera: Chironomidae). - Tijdschrift voor Entomologie 133: 149-195, figs.
1-112, tabs. 1-3. [ISSN 0040-7496]. Published 14 December 1990.
Fiftythree species of Chironomidae are recorded for the first time from rice fields of West
Bengal, India; 24 of these, including 5 new species, are new to India. Cryptochironomus
judicius sp. n. is described from male imago, pupa and larva; Harnischia tenuitubercula sp.
n., Paracladopelma aratra sp. n., P. sacculifera sp. n. and Polypedilum circulum sp. n. from
male imagines. The male imago of Stenochtronomus longipalpis (Kieffer) and all stages and
both sexes of Stictochironomus obscurus (Guha & Chaudhuri) comb. n., Clinotanypus
fuscosignatus (Kieffer), and Procladius noctivagus (Kieffer) are redescribed. Keys for the
identification of genera and species treated are presented for larvae, pupae and adults. A
general account of biology, emergence patterns and sex ratios concludes this paper.
Correspondence: Dr. P. K. Chaudhuri, Dep. of Zoology, University of Burdwan, Burdwan
- 713104, West Bengal, India.
Key words. - Chironomidae, rice fields, new species, keys, emergence, sex ratio.
Although rice fields have long been recognised as
a pre-eminent habitat for chironomids, there are
only few works dealing with this habitat. Thiene-
mann (1954) listed the species of midges from rice
fields of Southern Sumatra and West Java where the
larvae were used as food of carp. The harmful ac-
tivities of larvae of Cricotopus trifasciatus Panzer to
the rice seedlings in France were discussed by Ris-
bec (1952). Darby (1962) made a list of 36 species
from rice fields and its vicinities of California
(U.S.A.) and regarded 30 of them as actually inhab-
iting the rice paddies. Jones (1968) reported Chzro-
nomus tepperi Skuse as a pest of rice in Australia.
Martin & Porter (1977) presented the biology of
this species in the laboratory. Bognar (1958) in-
cluded chironomids in the arthropod pest group of
Hungary. Many chironomid species were reported
to injure rice seedlings in Japan (Tokunaga & Kur-
oda 1935, 1936, Yokogi & Ueno 1971, Ishihara
1972). Hashimoto et al. (1981) gave a brief descrip-
tion of the adult morphology of 32 species of chi-
ronomids in the rice fields of Thailand. Sasa &
Kikuchi (1986) made an account of 34 species from
the rice fields of Japan and stated that at least 40
species of chironomids were found to be breeding
in the rice paddy areas of Japan. There has been no
previous attempt to give an account of chironomids
of rice fields of the Indian subcontinent.
The present paper records 53 chironomid species
from the rice paddies of West Bengal, India. Of
these, 24 species, including 5 new species, are new
for India. A brief account of the biology, emergence
patterns and sex ratios is also presented.
MATERIAL AND METHODS
Adults were collected by sweeping in the rice
fields with a long handled insect net. Larvae were
collected from bottom mud in the rice fields with
the help of mud scrapers and a scoop sampler.
Larvae were reared in petridishes in the laboratory
following the procedure given by Chaudhuri et al.
(1983). Unless otherwise mentioned, all specimens
listed under ‘Material examined’ were collected in
West Bengal by S. Chattopadhyay.
The phenol-balsam technique, as used by Das
Gupta & Wirth (1968) was followed for microslide
preparation of larvae, pupae and adults. Larval and
pupal exuviae were also mounted on microslides in
Gum-chloral solution (Sasa 1978) without dehydra-
tion. Morphological terminology essentially fol-
lows Saether (1980).
Measurements are given in millimeters (mm);
the mean value is followed by the range and sample
size (n) in parentheses.
The types, paratypes and other specimens of this
study are provisionally kept in the insect collection
of the Entomology Laboratory, Department of
Zoology, University of Burdwan (B.U. Ent.), West
Bengal, India, but will be deposited in the National
Zoological Collections, Calcutta (NZC); National
Pusa Collections, New Delhi; British Museum
149
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
(Natural History), London (BMNH); United
States National Museum, Washington D.C.
(USNM) and Zoologische Staatssammlung, Mün-
chen (FRG) in due course.
LIST OF CHIRONOMID SPECIES TREATED
Subfamily: Chironominae
Tribe: Chironomini
1. Chironomus circumdatus (Kieffer)
2. Chironomus filitarsis Kieffer
3. Chironomus javanus Kieffer
4. Chironomus nudipes Kieffer
5. Chironomus samoensis Edwards
6. Chironomus striatipennis Kieffer
7. Cryptochironomus fulvus (Johannsen)
8. Cryptochironomus judicius sp. n.
9. Cryptochironomus rostratus (Kieffer)
10. Cryptochironomus subovatus Freeman
11. Dicrotendipes pelochloris (Kieffer)
12. Dicrotendipes septemmaculatus (Becker)
13. Endochironomus pekanus (Kieffer)
14. Harnischia acuta (Goetghebuer)
15. Harnischia incidata Townes
16. Harnischia tenuitubercula sp. n.
17. Harnischia viridula (Linnaeus)
18. Kiefferulus barbatitarsis (Kieffer)
19. Kiefferulus calligaster (Kieffer)
20. Microchironomus fuscitarsus (Guha & Chaudhuri)
21. Microchironomus tener (Kieffer)
22. Paracladopelma aratra sp.n.
23. Paracladopelma sacculifera sp.n.
24. Pentapedilum robusticeps Guha & Chaudhuri
25. Pentapedilum uncinatum (Goetghebuer)
26. Polypedilum aegyptium Kieffer
27. Polypedilum annulatipes (Kieffer)
28. Polypedilum ascium Chaudhuri, Guha & Das Gupta
29. Polypedilum chaudhurrii Chaudhuri, Guha & Das
Gupta
30. Polypedilum circulum sp. n.
31. Polypedilum lineatum Chaudhuri, Guha & Das Gupta
32. Polypedilum nubifer (Skuse)
33. Polypedilum medivittatum Tokunaga
34. Polypedilum suturalis (Johannsen)
35. Polypedilum yapensis Tokunaga
36. Stenochironomus hilaris (Walker)
37. Stenochironomus longipalpis (Kieffer)
38. Stictochironomus affinis (Johannsen)
39. Stictochironomus obscurus (Guha & Chaudhuri)
comb.n.
40. Kenochironomus flaviventris (Kieffer)
Tribe: Tanytarsini
41. Cladotanytarsus conversus (Johannsen)
42. Cladotanytarsus gloveri Ghosh & Chaudhuri
43. Cladotanytarsus multispinulus Guha, Das, Chaudhuri
& Choudhuri
44. Tanytarsus bifurcus Freeman
45. Tanytarsus commoni Glover
46. Tanytarsus fuscimarginalis Chaudhuri,
Ghosh
47. Tanytarsus vinculus Chaudhuri, Guha & Ghosh
Guha &
Subfamily: Tanypodinae
48. Clinotanypus fuscosignatus (Kieffer)
49. Procladius noctivagus (Kieffer)
150
50. Tanypus bilobatus (Kieffer)
51. Tanypus grandis Chaudhuri, Das & Debnath
52. Tanypus lucidus Chaudhuri, Das & Debnath
53. Tanypus tenebrosus Chaudhuri, Das & Debnath
KEYS TO GENERA AND SPECIES
Larvae
Key 1. Chironomini - genera
IEA ner naNOISe mentele
En ee Stictochironomus Kieffer (key 4)
SIVANTENDIDESCLMENEARERR ee oe ee 2
2. Mentum with broad, pale untoothed central region
dro Da: Cryptochironomus Kieffer (species 7-10)
- Mentum without broad, pale untoothed central re-
TOI O N n 3
3 Vental mes est 6 the se tee aa co 4
I Ventralktubulusfabsenter ee 5
4. Segment XI with 2 pairs of ventral tubules........
Pale GLIO Chironomus Meigen (species 1-6)
- Segment XI with 1 pair of ventral tubules....... 6
5. SIblade-like. Mentum with a trifid median tooth and
Glpairsiomlateralitee he
Res A A ta Microchironomus Kieffer (key 3)
- SI plumose. Mentum with a bifid median tooth and
VApa1rstof@lateraliteeth Eee ieee eee
ELITE Polypedilum Kieffer (species 26-35)
62 BectenlepipharyngisiwithDäteecher we RE E ene
Re Dicrotendipes Kieffer (species 11, 12)
— Pecten epipharyngis with 16-19 teeth............
RITIRO. SN Kiefferulus Goetghebuer (key 2)
pa! 2. Kiefferulus Goetghebuer
SI pectinate. Ventromental plate with wrinkled outer
surface. Posterior parapods each with 15-16 well-
Selerotisedhelawst. GE Ge LE OOMEN ee
SERRE PIER I 18. barbatitarsis (Kieffer)
- SI plumose. Ventromental plate with smooth outer
surface. Posterior parapods each with 13-14
Cla WS) ede 19. calligaster (Kieffer)
Key 3. Microchironomus Kieffer
1. Mandible with pointed lateral teeth; seta interna with
3 main plumose branches; pecten mandibularis well
developed. Ventromental plate wrinkled at its outer
PRR pia wo ABS 20. fuscitarsus (Guha & Chaudhuri)
— Mandible with flat lateral teeth; seta interna with 3-
4 filaments; pecten madibularis absent. Ventromen-
tal plate smooth at its outer margin...... 21. tener
(Kieffer)
Key 4. Stictochironomus Kieffer
1. SITandS I leaf like; pecten epipharyngis comprised
of 3 toothed plates, median one with 3 and each
lateralfoneswithtSntee ne EEE
hin E Hen 38. obscurus (Guha & Chaudhuri)
— Sand SII plumose; pecten epipharyngis comprised
of 3 toothed plates, median one with 3-5 and each
lreralonewithpitee he ee
Key 5. Tanypodinae - genera
1. M appendage without pseudoradula.............
IT Gon Tanypus Meigen (species 50- 22
— M appendage with pseudoradula.............. 2
ho
Dorsomental teeth not located on distinct plate.
Mandible strongly hooked with large, pointed basal
tooth eulanwithgONeeCh EEEN een:
Nen Clinotanypus Kieffer (species 48)
— Dorsomental teeth located on distinctly defined plate.
Mandible not as above. Ligula with 5 teeth........
Fae tee aA Procladius Skuse (species 49)
Pupae
Key 6. Chironomini - genera
l. Anal segment dorsally with forked, posteromedian
PLOCESS ANNEE PA PE Ae en. kesh eeen
NE VA Cryptochironomus Kieffer (species 7- 19
- Anal point without such process ..............
Segment VIII with 2 pair of simple B
Spürsyendin pain apo In EEEN EE 3
- Segment VIII with 1 pair of caudolateral spurs, each
withemorenthanslipoin NEEN STE 4
Thoracic horn with a bunch of finely branched fila-
ments ee or SRO o OR n
MR ATTO Dicrotendipes Kieffer (species 11, 12)
- Thoracic horn with 4 main bunches of finely
branchedifilaments er as AE
N IKT TI Microchironomus Kieffer (key 8)
4. Segment VI with only 3 pairs of filamentous lateral
ho
Wo
SAE nd LE ech aren A 5
- Segment VI with more than 3 pairs of filamentous
lateralisetacy RE LEUR MOED TINS EURE 6
54 Erontal setae short and slenden. an MAANEN
MEETS FI RENT Polypedilum Kieffer (species 26-35)
- Frontal setae long, robust and broad.............
RE A EIER Stictochironomus Kieffer (key 9)
6. Tergite VII without median shagreen............
EEL) Kiefferulus Goetghebuer (key 7)
- Tergite VII with median shagreen...............
ste enne Chironomus Meigen (species 1-6)
el 7. Kiefferulus Goetghebuer
Thoracic horn with a bunch of fine filaments. Seg-
ment I with 1 pair of lateral setae, dorsal setae ab-
SCM CE teaches ite BAL 18. BR (Kieffer)
— Thoracic horn with 2 plumose branches. Segment I
without lateral setae, dorsal setae 2 pairs..........
RN ne RANG Nb LL LE 2 19. calligaster (Kieffer)
Key 8. Microchironomus Kieffer
1. Frontal seta absent. Tergite VII bare. Segments II-
VII with 1, 2, 2, 3, 4, 4, 2 pairs of dorsal setae.
Segments V- VI without filamentous lateral setae
a2 20. fuscitarsus (Guha & Chaudhuri)
— Frontal seta present. Tergite VII with subbasal trans-
verse patches of shagreen. Segments II-VIII with 3,
6, 5, 4, 6, 6, 4 pairs of dorsal setae. Segments V-VI
withitilamentoustalteralisetae EE
EET EN eren (ie)
Key 9. Stictochironomus Kieffer
1. Thoracic horn with a bunch of finely branched fila-
ments, segment I without dorsal setae............
erde And ee 36. obscurus (Guha & Chaudhuri)
- Thoracic horn with 2 main bunches of finely
branched filaments, segment I with dorsal setae
enten a QUE Ue EOE Sheer 37. affinis (Johannsen)
Key 10. Tanypodinae - genera
1. Thoracic horn without plastron plate............
TRA a Tanypus Meigen (species 50-53)
CHAUDHURI & CHATTOPADHYAY: Chironomids
— Thoracic horn with plastron plate............. 2
2. Plastron plate reduced. Anal fin with a fringe of
SPINEsOnspinUle WEEKE IN
Mr tet EN Procladius Skuse (species 49)
— Plastron plate broad. Anal fin without a fringe of
SITES Or GHIMMNES', solange ago I Aedo og cor
Adults
Key 11. Chironomini - genera
1. Anal point with basolateral setaceous lobe........
ae aceto Microchironomus Kieffer (key 17)
— Anal point without basolateral setaceous lobe.... 2
2MPosteniontibiawithonespu 2. Sn are. 3
Posterior tibia with two spurs ................ 5
3. Macrotrichia present on wing membrane, wing with-
OUUSPOLSOLIMALKIDE SEE IN NI
MATITE: Pentapedilum Kieffer (key 19)
— Macrotrichia absent on wing membrane, wing with
Or without spotsior markings) EE PET ET EN CEE 4
4. Mosonotum with a distinct tubercle at the posterior
end of the median vittae. Femora and tibiae usually
PIN CALAN bes LANDEN PANIER A
REATO ee ae Stictochtronomus Kieffer (key 22)
— Mesonotum without such tubercle at the posterior
end of median vittae. Femora and tibiae not distinctly
INGE Mer rr en ACIREALE
Fey eA Ue Aca Polypedilum Kieffer (key 20)
5. Inferior volsella reaching well beyond tip of gonoc-
oxite and bearing long curved setae............
— Inferior vosella not reaching beyond tip of gonocoxite
and without long curved setae............... 11
6. Inferior volsella extremely broad and bulbous api-
CAGE PAR ARS TRE EEE ET
nni pari dans Kiefferulus Goetghebuer (key 16)
- Inferior volsella slender, not or scarcely enlarged dis-
NIC REN NEE REITER 7
7. Inferior volsella strongly bowed dorsoventrally with
an expanded clubbed or slightly to deeply bifid to
trifid apex bearing many strong setae............
EN RE LINO Dicrotendipes Kieffer (key ee
— Inferior volsella not as above.................
8. Anal point very broad. Superior volsella short, er
Andy pubeSCeNt tee TOA I IT
Ba: paren RECHT Xenochironomus Kieffer (species 40)
— Anal point narrow. Superior volsella not as
CNR RR A n TENEN 9)
9. Superior volsella strongly chitinized and bare except
for a few long basal setae, usually ending a spur or
NOOK Petree ee rece hee er lee tat Ren TI ENI
in Se Ed Chironomus Meigen (key ne
- Superior volsella not as above ...............
10. Antepronotum extending to anterior edge of scutum
but deeply divided into two anterior lobes. Inferior
volsella slender bearing 2 long apical setae and with-
OUIICEHINIDA NS pIN CRANE ree tere
RE TATE Endochironomus Kieffer (species 13)
- Antepronotum not as above. Inferior volsella very
long bearing no apical setae but with a well differen-
tiatediterminalus Pine enn BROS CR OR ACRI
Ee Stenochironomus Kieffer (key 21)
11. Superior volsella short and broad, densely pubescent
with a few long setae. Inferior volsella in the form of
aismallipubescentipad ARR
RER Paracladopelma Harnisch (key 18)
— Both volsellae not as above ................. 12
151
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Gonostylus short and broad. Superior volsella short,
bsoadtandipubescen re OI
re. Cryptochironomus Kieffer (key 13)
Gonostylus longer. Superior volsella reduced, occa-
stonallysabsenten Hull ite ae SE
Harnischia Kieffer (key 15)
Key 12. Chironomus Meigen
Ils
D |
(US)
Wing with spots or markings..................
EN a ati 6. striatipennis Kieffer
Wing without spots or markings.............. 2
Antepronotals 2. Scutellum with 8 setae..........
Ar AALTEN 3. javanus Kieffer
Antepronotals absent. Scutellum with more than 17
SCHAREN EA EN Eon 3
Corona with 2 setae. Gonostylus oblong with blunt
IPER IR OEE 4. nudipes Kieffer
Corona bare. Gonostylus not as above .......... 4
Branchiolum with 3 setae. Haltere setose.........
RARE ANR ARE EE A OE 5. samoensis Kieffer
Branchiolum with 2 setae. Haltere bare......... 5
Tergites II-V with grey, oval median spots........
1. circumdatus (Kieffer)
Tergites I-IV with brown, square median spots 2.
filitarsis Kieffer
Key 13. Cryptochironomus Kieffer (males)
IL
2
Seutellum)wichiO=tOlsetae Ate sees PRE 2
Seutellumy with) -26)setaetnes 94. see 3
Frontal tubercles prominent and each with an apical
knob. Superior volsella more or less globular, inferior
volsella small and superposed on superior vol-
Sellar Aen: 7. fulvus (Johannsen)
Frontal tubercles small, rounded and each without an
apical knob. Superior volsella short, digitiform; infe-
Bor WOKE: PSE NCR RR RR N.
10. subovatus Freeman
Haltere light yellow with 11-13 setae. Superior vol-
sella cone shaped bearing 2 setea, inferior volsella
ABSENCE IDE EEE Ae 8. judicius sp. n.
Haltere yellowish green and bare. Superior volsella
thumblike bearing 3 apical setae, inferior volsella
LES REE ere 9. rostratus (Kieffer)
Key 14. Dicrotendipes Kieffer (males)
1.
Scutellum with 6-8 setae in one transverse row. Wing
without spots. Inferior volsella gently bowed ven-
RUN RS eat 11. pelocholoris (Kieffer)
Scutellum with 12 setae in two transverse rows. Wing
with 6 grey spots. Inferior volsella bifurcated into 2
Bluntylobe sag saris. LA RD
Key 15. Harnischia Kieffer (males)
Il
Male hypopygium without well marked gonocoxite
AAIPONOS VUS EN ene REE
17. viridula (Linnaeus)
Male hypopygium with well marked gonocoxite and
CONOSCIUTE OI 2
Élaltere bare Res 16. tenuitubercula sp. n.
ÉTAITÉTESS ET OS CPP EPP PRE PER 3
Brachiolum with 2 setae. Superior volsella bearing 3
long apical setae, inferior volsella rudimentary with-
QUEON LISCIO
14. acuta (Goetghebuer)
Brachiolum with 1 seta. Superior volsella bearing 1
apical seta, inferior volsella absent..............
15. meidata Townes
Key 16. Kiefferulus Goetghebuer (males)
il,
Scutellum with 20 setae in two transverse rows. Bra-
chiolum with 4 setae. Anal point well developed,
toneue KE WAAN NN EREN ANNE
AREA LUNES I N CEES? 18. barbatitarsis (Kieffer)
Scutellum with 6 setae in one transverse row. Brachi-
olum with 2 setae. Anal point dark, more or less peg-
TRE RE re EREN AE 19. calligaster (Kieffer)
Key 17. Microchironomus Kieffer (males)
I
Anal point with a small setaceous basolateral lobe.
Gonostylus with 15-17 setae at its inner margin. Su-
perior volsella little curved with 1 apical seta . .....
Eh deken 20. fuscitarus (Guha & Chaudhuri)
Anal point with a large setaceous basolateral lobe.
Gonostylus with 8-10 setae at its inner margin. Su-
perior volsella long and slender with 2-3 apical
SPINE NEE 21. tener (Kieffer)
Key 18. Paracladopelma Harnisch (males)
IL
Frontal tubercles absent. Prescutellar present. Spurs
of hind tibia equal. Anal point tubular 22. aratra
sp. n.
Frontal tubercles present. Prescutellar absent. Spurs
of hind tibia unequal. Anal point with saccular
CD PEI AEEA 2 23. sacculifera sp. n.
Bey 19. Pentapedilum Kieffer
Scutellum with 26 setae in two transverse rows. Bra-
chiolum with 7 setae. Anal point robust, superior
volsellatwithountalatera rer
ana un 24. robusticeps Guha & Chaudhuri
Scutellum with 10-11 setae in a transverse row. Bra-
chiolum with 1 seta. Anal point rod like, rather
pointed at tip, superior volsella with 1 lateral
Sete nid aero DAN 25. uncinatum (Goetghebuer)
Sp 20. Polypedilum Kieffer (males)
Bi wl nie
Wing with spots or markings................ 2
Wing without spots or markings.............. 6
ARslessichannl ine 31. lineatum Chaudhuri et o
ARgmore;thanyliyp:{otha bian RME
Frontal tubercles present ...... 32. nubifer ase
Erontalitubercles absente se
Anal point narrow with pointed apex............
SARAI VIRA FAR voli RR ANT, 27. annulatipes (Kieffer)
Anal point broad without pointed apex.........
Prealars 3. Scutellum with a row of setae.........
SSR RO ACI EN LIRA 26. aegyptium Kieffer
Prealars 6. Scutellum with 2 rows of setae.........
................ 29. chaudhuru Chaudhuri et al.
Hlaltere’setosen RR RE FORI SCAN 7
Haltereibarer: mlMet Mee RL EE 8
Superior volsella curved with blunt tip and 1 long
lateral¥setasnearithe base 4 ane E
IRR e SIOE nio tse 30. circulum sp. n.
Superior volsella scimitar-like without lateral......
SECRET II 33. medivittatum Tokunaga
Superior volsella without lateral seta.............
MR aa 28. ascıum Chaudhuri et al.
Superior volsella with lateral seta............. 9
Prescutellar present, scutellum with 6 setae. Tergites
I-VII with brown median band.................
Rai 34. suturalis (Johannsen)
Prescutellar absent, scutellum with 4 setae. Tergites
woitormly brow lease eee eae eee
Key 21. Stenochironomus Kieffer (males)
I. Scutellum with 30 setae in two transverse rows. Wing
with pale median and apical bands ..............
SIDE cet US Leone oe ola a Oe 36. hilaris (Walker)
— Scutellum with 14 setae in one transverse row. Wing
UNMALRE nrg aoe 37. longipalpis (Kieffer)
Key 22. Stictochironomus Kieffer (males)
l. Corona with 4 setae. Anal point with an apical
KID LEI 39. obscurus (Guha & Chaudhuri)
— Corona bare. Anal point without apical knob
Va eae PARI O LA A 38. affinis (Johannsen)
Key 23. Tanytarsini - genera
l. Median volsella long and branched..............
TE NE Cladotanytarsus Kieffer (key 24)
— Median volsella short and brush-like .............
DOO RO ANE ats ee Tanytarsus v. d. Wulp (key 25)
Key 24. Cladotanytarsus Kieffer (males)
[eeekcontalitubercles;absent spawn re
SEIT, oe 43. multispinulus Guha et al.
SME contalituberclesypresent amine EE saat 2
Anal point inverted ‘bell-shaped’ and with 6-8 punc-
TURES ere RT centre 42. gloveri Ghosh & Chaudhuri
- Anal point short without punctures..............
sie PAS aia CAE nae 41. conversus (Johannsen)
ho
Key 25. Tanytarsus v. d. Wulp (males)
1. Anal point bifurcated ........ 44. bifurcus Freeman
POLS ID ERE RE O 2
DEB Tacho lu maw hi ASTA CRE SITI
A ORA 46. fuscimarginalis Chaudhuri et al.
See branchiolumbwithwlesetal sn en er er 3
Scutellum with 6 setae. Anal point with 4 lateral setae
andiaysubapicaluovalilobey u... 2.20. ET ee
ed cn eee Bee raat eae unt ee 45. commont Glover
- Scutellum with 4 setae. Anal point without lateral
setae andisubapiealtovalilobem EPP E CCE EEE
Pyare ci AR 47. vinculus Chaudhuri et al.
a 26. Tanypodinae - genera
Fourth tarsal segment bilobed and shorter than
fifteen eee eee Clinotanypus Kieffer (species 48)
- Fourth tarsal segment cyclindrical and not shorter
thangtı ches ea ate label 2
N
Distance between FCu and MCu less than one third
these nh dC RE O
E ROSE EA Tanypus Meigen (key 27)
- Distance between FCu and MCu at least half the
ERETTO ACU
Ds 27. Tanypus Meigen (males)
Brachrolunawjlthelase ta ee 2
= Bradmolmin yay? Ganesan does ved eedas adds 3
Scutellum with 10 large setae at the posterior border
and 20-22 small setae arranged in the anterior …
©
LEONE 52. lucidus Chaudhuri et al.
— Scutellum with 6-10 setae at the posterior
CLIO EEE 50. bilobatus (Kieffer)
Wo
Tarsomeres I-IV with apical dark brown bands, tar-
SOMERE AVA DLOW DEE
kel etn nti tienes, Aes 51. grandis Chaudhuri et al.
- Tarsomeres uniformly brown..............-...
NS: 53. tenebrosus Chaudhuri et al.
CHAUDHURI & CHATTOPADHYAY: Chironomids
SYSTEMATIC ACCOUNT
l. Chironomus circumdatus (Kieffer)
Tendipes circumdatus Kieffer, 1916: 110.
Chironomus circumdatus, Hashimoto 1977: 83; Sasa
1978; 11; Hashimoto et al. 1981: 5; Sasa & Hasegawa
1983: 316; Sasa & Kikuchi 1986: 18.
Material examined. - 11 4, Howrah, 4 September 1987;
6 8, Haldia, 16 October 1987; 4 6, Burdwan, 7 September
1987.
Distribution. — Japan, Korea, Taiwan, Thailand
and India (present record).
Differential diagnosis. — i) scutellum with 24-26
setae, ii) abdominal tergites I-V each with grey oval
median spot, iii) a deep constriction between seg-
ments VII and VIII and iv) hypopygium with stout
anal point and superior volsella slightly curved.
2. Chironomus filitarsis Kieffer
Chironomus filitarsis Kieffer, 1911: 160; Chaudhuri &
Guha 1987: 27.
Material examined. - 6 &, Burdwan, 30 May 1986.
Distribution. — India.
Differential diagnosis. — i) scutellum with 22-24
setae, ii) fore tibial scale blunt, bearing 2 long setae,
iii) tergites II-IV each with square shaped dark
brown spot and iv) superior volsella bent inward
with pointed apex.
3. Chironomus javanus Kieffer
Chironomus javanus Kieffer, 1924b: 263; Johannsen
1932: 536; Tokunaga 1964: 566; Hashimoto et al. 1981:
7; Sasa & Hasegawa 1983: 317; Sasa & Kikuchi 1986:
18.
Material examined. - 6 &, Farakka, 29 September 1986;
9 &, Howrah, 4 September 1987.
Distribution. — Caroline Islands, Japan, Java,
Marshall Islands, Thailand and India (present re-
cord).
Differential diagnosis. — i) antepronotum bear-
ing 2 antepronotals, ii) supraalars absent, iii) fore
tibial scale bearing 4 long setae, iv) tarsomeres I-III
brown at apex, tarsomeres IV-V dark brown and v)
hypopygium with long and tubular anal point, go-
nostylus with swollen basal half and superior vol-
sella curved.
Remarks. - The specimens collected from the
rice fields conform fully with those described by
Hashimoto et al. (1981) and Sasa & Hasegawa
(1983), except minor morphometrical differences.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
4. Chironomus nudipes Kieffer
Chironomus nudipes Kieffer, 1911: 164; Chaudhuri &
Guha 1987: 27.
Material examined. - 6 8, Uttarpara, 9 September 1987.
Distribution. - India.
Differential diagnosis. - ı) fore tibial scale with
2 long setae, ii) tarsomeres III-V brown, iii) tergites
VI-VII each with dark brown median spot and iv)
hypopygium with stout, arrow shaped anal point,
gonostylus oblong with blunt apex, superior vol-
sella bow-shaped and inferior volsella curved.
5. Chironomus samoensis Edwards
Chironomus samoensis Edwards, 1928: 67, Tokunaga
1964: 567, Hashimoto 1977: 82, Sasa & Hasegawa
1983: 316; Sasa & Kikuchi 1986: 18.
Chironomus dorsalis Tokunaga, 1940a: 220 (see Tokunaga
1964).
Chtronomus eximius Johannsen, 1946: 193 (see Tokunaga
1964).
Chironomus flaviplumus Tokunaga, 1940b: 294; Sasa
1978: 12; Ree & Kim 1981: 136.
Material examined. — 8 larvae, Berhampur, 2 January
1988; 4 larvae, 12 6 and 12 9 pupae, 5 6 and 7 9 exuviae,
10 6 and 10 Q, all reared.
Distribution. - Caroline Islands, Japan, Java, Ko-
rea, Marshall Islands, Samoa, S. Mariana Islands
and India (present record).
Differential diagnosis. - Larva: i) sickle shaped S
I, ii) pecten epipharyngis with 14 teeth, iii) pre-
mandible with 2 unequal apical teeth, premandib-
ular brush short, iv) mandible with 2 seta externa,
v) bladelike anterior chaeta of maxilla and vi) Xth
abdominal segment with apically pointed caudolat-
eral processes.
Pupa: i) cephalic tubercles conical, bearing sub-
apical setae, ii) thoracic horn with a bunch of finely
branched filaments, iii) pedes spurii A present
caudolaterally on segment IV-VII, iv) segment I-
VIII with 4, 5, 3, 4, 3, 4, 6, 2 pairs of dorsal setae
and 1, 3, 3, 3, 4, 4, 4, 5 pairs of lateral setae, v)
caudolateral spur on segment VIII with 2-4 unequal
points or branches, vi) G/F 0.66 in male and 0.76
in female.
Adult: i) frontal tubercles well developed, ii)
prescutellar present, scutellars in two transverse
row, iii) brachiolum with 3 setae, iv) RM and FCu
almost in the same line, v) colour pattern of legs,
vi) tergites II-VI with grey median spots in male,
vil) superior volsella of male hypopygium shoe
shaped, inferior volsella with 13-15 apically bifid or
trifid setae and viii) genitalia of female with prom-
inent apodeme lobe.
154
6. Chironomus striatipennis Kieffer
Chironomus striatipennis Kieffer, 1910: 236; Chaudhuri
& Guha 1987: 27.
Material examined. - 6 @, Palasi, 18 August 1986; 3 8,
Uttarpara, 24 February 1987; 8 6, Burdwan, 6 September
1987.
Distribution. — India and Taiwan.
Differential diagnosis. — 1) scutellum with 14-18
setae, ii) markings in wings, iii) tarsomeres I-III
dark brown at apices, iv) tergites II-V with brown
median spots and v) hypopygium with bent anal
point and curved superior volsella.
7. Cryptochironomus fulvus (Johannsen)
Chironomus fulvus Johannsen, 1905: 224.
Chironomus (Cryptochironomus) fulvus, Johannsen
1057259)
Tendipes (Cryptochironomus) fulvus, Darby 1962: 50.
Cryptochironomus fulvus (Johannsen); Townes 1945: 98;
Hashimoto et al. 1981: 16; Sasa & Hasegawa 1983:
322; Sasa & Kikuchi 1986: 20.
Material examined. - 5 8, Berhampur, 17 August, 1986.
Distribution. - Japan, Thailand, U.S.A. and India
(present record).
Differential diagnosis. — i) frontal tubercles
prominent and each with an apical knob, ii) ante-
pronotum reduced, iii) scutellum with 10 irregular
setae, iv) gonocoxite and gonostylus ankylosed, v)
superior volsella semiglobular, finely setose and
with 3 long setae and vi) inferior volsella small,
superposed on superior volsella bearing one apical
seta.
8. Cryptochironomus judicius sp. n.
Type material. - Holotype 4, India, West Bengal, Ut-
tarpara, 9 September 1987, leg. S. Chattopadhyay (Type
no. 182 B.U. Ent.). Paratypes: 4 8, data as holotype; 5 8,
8 pupae and 8 exuviae, 6 larvae and 4 exuviae, all reared
in laboratory.
Differential diagnosis. - Larva: i) AR 0.96, ii) S
I and S II blade like, iii) premandible with 4 unequal
apical teeth, premandibular brush dense, short, iv)
pecten mandibularis absent, v) each lateral comb of
mentum with 7 dark pointed teeth; 7th tooth min-
ute, vi) maxilla with 2 sensilla basiconica and vii)
procercus with 8 anal setae.
Pupa: i) cephalic tubercles elevated and bearing
subapical setae, ii) thoracic horn with an oval base
and finely branched filaments, iii) pedes spurii A
caudolateral on segment IV and pedes spurii B bas-
olateral on segment I, iv) tergite I bare, v) lateral
setae on segments VI-VIII filamentous and vi)
caudolateral spur on segment VIII with 4-5 stout
spines.
Adult: i) frontal tubercles very small, ii) scutel-
lum with 26 irregular setae, iii) haltere light yellow
with 11-13 setae, iv) fore tibia with a blunt scale
bearing 3 long setae, v) colour pattern of terso-
meres, vi) pulvilli moderately developed, vii) ter-
gites II-VI with grey caudal spots, viiii) gonostylus
dark brown and iv) superior volsella cone shaped
bearing 2 setae.
Remarks. - The species is named judzc14s for its
pointed teeth of the lateral combs of the larval
mentum. The present species resembles C. neont-
licola Kieffer as described by McLachlan (1969) in
respect to the larval antennal characteristics. It
shares similarities in the mandible with C. /indneri
Freeman, C. diceras Kieffer and C. conus Mason,
1985, in the ventromental plate with C. ramus Ma-
son, 1985 and in mentum and maxilla with C. emi-
nentia Mason. The pupal respiratory organ of C.
redekei (Kruseman) appears to be similar to that of
the present species. The adults of the new species
look like C. conus Mason in the shape and structure
of the anal point of the male hypopygium.
Description
Adult. - Male: Body 3.98 (3.91-4.13, n= 9) long,
wing 1.47 (1.44-1.48, n = 10) long and 0.46 (0.45-
0.48, n = 10) wide.
Head: Brown. Vertex with 8-11 setae (Iv 2-3, OV
5-6, PO 1-2). Corona bare. Clypeus with 23-25 setae,
clypeal ratio 1:1. Maxillary palp pale brown, ratio
of palpomere length I-V: 9 : 14 : 38 : 33 : 52; L/W.
4.75. Frontal tubercles (fig. 1) small. Antenna
brown, ratio of flagellomere length I-XI: 17 :7 :8:
6:6:626:6:6:5 : 254; AR 3.36; pedicel ratio
OAS CARO AER NED
Thorax: Yellow: Antepronotum collar-like with
slight emargination in the middle, antepronotals O.
Mesonotum with 3 brown vittae. Acrostichals 24-
25 irregularly biserial, dorsocentrals 17 uniserial,
humerals 5, prealars 5. Scutellum with 26 irregular
setae, postscutellum dark brown and bare. Wing
(fig. 2): Hyaline. Brachiolum with 2 setae and 19-
21 sensilla campaniformia. R with 24, R, with 11-
12 and R,;; with 26-28 setae; R,4; meets C at a
distance of 0.14 from R,; RM and FCu almost in
same line. An ends below FCu. Squama with 12-13
setae. Haltere light yellow with 11-13 setae. CR
0.89; VR 1.12.
Legs: Yellow. Fore tibia with a blunt scale (fig. 3)
bearing 3 long setae. Spurs of mid tibia subequal,
0.021 and 0.018 long, ratio of length of spurs to the
apical diameter of mid tibia 7 : 15 and 6 : 15; spurs
of hind tibia unequal 0.024 and 0.018 long, ratio of
length of spurs to the apical diameter of hind tibia
8 : 18 and 6 : 18. Apex of tarsomeres I-III, tarso-
meres IV-V of fore leg and tarsomere V of mid and
hind legs brown, sensilla chaetica indistinguishable.
Pulvilli moderately developed. Proportions and ra-
tios of leg-segments in table 1.
CHAUDHURI & CHATTOPADHYAY: Chironomids
Abdomen: Yellow. Tergites II-VI (fig. 4) with
grey caudal spots. Hypopygium (fig. 5) with slender
anal point 0.057 long bearing 7-8 bosolateral setae.
Gonocoxite with 5-6 setae; gonostylus stout, dark
and with 8-9 setae at its inner margin and 8-10 setae
over it. Superior volsella cone shaped bearing 2
setae; inferior volsella absent. Transverse sternap-
odeme 0.033, lateral sternapodeme 0.147, coxapo-
deme 0.057 and phallapodeme 0.12 long. HR 0.39;
EINMIE89)
Female. - Unknown.
Pupa. - Reddish brown. Exuviae pale brown.
Body 5.19 (5.17-5.24, n = 8) long in male. Female
pupa and exuviae unknown.
Cephalothorax: Brown. Frontal apotome (fig. 6)
with elevated cephalic tubercles, 0.02 long and
0.048 diameter at base, subapical faint frontal seta
0.027 long. Antennal sheath in 8 (fig. 7) 0.72 long.
Thorax rugose; wing sheath 0.99 long; Thoracic
horn (fig. 8) with oval base 0.051 wide and a bunch
of finely branched filaments. 2 pairs of precorneal
setae.
Abdomen (fig. 9): Pale brown. Pedes spurii À
caudolateral on segment IV, pedes spurii B basolat-
eral on segment I. Tergite 1 bare; tergite II (fig. 10)
with median patch of shagreen and with a caudal
row of numerous hooklets; tergites III-IV with sub-
basal transverse patch of shagreen; tergites V-VI
with shagreen in the form of inverted ‘T’; tergite
VII with median longitudinal and tergite VIII with
subbasal patch of shagreen. Segments II-VIII with
3, 4, 4, 4, 3, 4, 2 pairs of dorsal setae and segments
II-VII with 2, 2, 2, 4, 4, 4, 5 pairs of lateral setae,
on segments VI-VIII filamentous; segment VIII
with 1 pair of caudolateral spur 0.024 long, each
with 4-5 stout spines. Anal fin (fig. 9) 0.31 long
with forked processes and numerous filamentous
setae. Genital sac in @ (fig. 11) 0.35 long. G/F 1.12
in d.
Fourth instar larva. - Colouration red, exuviae
white, head capsule, claws of anterior and posterior
parapods, and anal setae brown. Body 7.97 (7.64-
8.09, n = 10) long.
Head: Brown. Occipital margin light brown.
Ventral head capsule (fig. 12) 0.35 long and 0.29
wide. Two eye spots connected to each other. An-
tenna (fig. 13) five-segmented, basal antennal seg-
ment 0.036 long and 0.013 wide, with a ring organ
0.006 in diameter, distance to ring organ from base
0.024; blade of basal antennal segment 0.03 long;
accessory blade 0.021 long; blade of antennal seg-
ment II 0.013 long; Lauterborn organ small, at apex
of antennal segment III; ratio of antennal segment
length EME SE GEMINI FARSO OG mEabral
lamella unmarked; S I (fig. 14) blade like 0.093 long,
between the bases of two SI 0.018 long; S II blade
155
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 1. Proportions and ratios of leg-segments of adult Chironomidae.
Ee Ti tal ta2 tai
8. Cryptochironomus judicius sp.n.
Fore 54 46 71 36 27
Mid 49 45 29 14 10
Hind 56 Dy] 45 21 19
16. Harnischia tenuitubercula sp.n.
Fore 27 24 49 28 20
Mid 33 29 7) 11 6
Hind 42 44 28 Il7 15
22. Paracladopelma aratra sp.n.
Fore 31 21 37 19 15
Mid 29 23 12 6 4
Hind 32 31 19 11 10
23. Paracladopelma sacculifera sp.n.
Fore 49 30 67 31 22
Mid 44 37 23 11 8
Hind DI 49 34 18 16
30. Polypedilum circulum sp.n.
Fore 65 34 73 46 29
Mid 58 53 31 15 11
Hind 68 58 45 24 22
39. Stictochironomus obscurum (Guha & Chaudhuri)
Fore 43 39 51 30 24
Mid 51 45 26 14 11
Hind 49 51 40 DI
48. Clinotanypus fuscosignatus (Kieffer)
Fore 72 85 69 35 22
Mid 75 77 53 22 12
Hind 68 87 57 28 20
49. Procladius noctivagus (Kieffer)
Fore 40 DI, 58 18 13
Mid 46 45 29 14 10
Hind 42 50 34 17 13
like; S III simple; S IV 3 segmented; 4 chaetae; 2
spinulae; 3-4 chaetulae laterales; chaetulae basales
absent. Pecten epipharyngis comprised of 3 minute
toothed plates. Premandible (fig. 15) 0.054 long
with 4 unequal teeth, premandibular brush dense
and short. Mandible (fig. 16) 0.084 long with 1
apical and 2 lateral teeth, seta subdentalis 0.015
long, seta interna with 4 long and 2 short filaments,
3 seta interna; pecten mandibularis absent; ring
organ 0.006 in diameter, distance to ring organ
from base 0.009; inner margin smooth. Maxilla
with poorly developed lamellae of galea; broad an-
terior chaeta; leaf like lacinial chaetae; antaxial seta
and paraxial seta developed, sensilla basiconica 2;
maxillary palp 3 segmented, firs segment of max-
illary palp 0.021 long, 0.027 wide with ring organ
0.003 in diameter and 2-3 sensilla at its apex; a few
chaetulae palpiger, 2 setae maxillaris, SM, and SM,
156
ta4 ta5 LR BV SV BR
21 11 1.54 1.80 1.69 2.14
6 0.64 gl 4.48 2.4
11 7 0.79 DD 3.05 227)
14 8 2.04 1.57 1117 2.80
4 3 0.59 2229) 4.77 3.20
8 5 0.64 255 3.07 4.43
10 6 1.76 1.78 1.68 25
3 255 0.52 4.13 5.47 2.8
5 4 0.61 2.56 Spill 22
17 8 255 1.87 1.68 2.28
5 4 0.62 DIA 4.76 3.43
9 6 0.69 275 222) 4.14
23 11 1.15) 1.58 1977 2.83
7 D 0.58 3.74 4.83 SLI
12 6 0.77 2.67 3.15 3.87
19 11 il 1.58 192 3A
8 6 0.58 Sallis 3.84 3.86
11 8 0.78 2.41 2.70 6.14
7 10 0.81 3.05 4.02 2.54
5 8 0.69 4.36 6.08 3.30
7 9 0.65 3311 4.30 2.95
9 7 0.72 279 211 2.14
7 6 0.64 3.24 3.96 3.14
8 6 0.68 2.86 SA 2.87
close to the palpiger, SM; and SM, close together
on the clerite of carido. Prementohypopharyngeal
complex (fig. 17) with prementum 0.018 wide, me-
dian lamella distinct, 2 well developed paramedian
lamellae, 3 of sensilla and 5 of chaetulae. Mentum
(fig. 18) 0.033 long and 0.072 wide with broad, pale
untoothed central region, flanked by oblique lateral
combs of 7 dark pointed teeth. Ventromental plate
(fig. 19) fan-shaped 0.03 wide, outer margin
smooth, with faint rays ending before the margin.
Abdomen: Brown. Procercus 0.028 long and
0.014 wide, each with 8 anal setae, 0.055 long and
2 short lateral setae. 2 supraanal setae, 0.18 long;
Sa/An 0.33. Anterior parapods with numerous
weakly sclerotised claws, posterior parapods (fig.
20) 0.182 long with 12-13 sclerotised, variable claws
(fig. 21). Anal tubules (fig. 20) conical 0.098 long.
9. Cryptochironomus rostratus (Kieffer)
Chironomus rostratus Kieffer, 1911: 164.
Chironomus (Cryptochironomus) rostratus; Goetghebuer
1928: 84.
Chironomus (Chironomus) rostratus; Edwards 1929: 390.
Tendipes (Cryptochtronomus) rostratus, Kruseman 1933:
187.
Cryptochironomus rostratus, Pinder 1978: 116; Ree &
Kim 1981: 143.
Material examined. - 6 4, Howrah, 4 September 1987.
Distribution. - Belgium, The Netherlands, Eng-
land, Germany, India and Korea.
Differential diagnosis. — 1) frontal tubercles
small, ii) haltere yellowish green and bare, iii) ap-
ical hooks of hind femora pale and inconspicuous,
iv) gonostylus short, thick and pointed apex, v)
superior volsella thumb-like bearing 3 apical setae
and vi) inferior volsella small, lobe-like densely
setose.
10. Cryptochironomus subovatus Freeman
Chironomus (Cryptochironomus) subovatus Freeman,
1954: 20.
Cryptochironomus subovatus; Freeman 1955a: 375; Free-
man 1957: 390; Freeman & Cranston 1980: 189.
Material examined. - 5 8, Berhampur, 17 August 1986;
Holotype & (Chir. 437.B), Cape Province Berg R., 27
March 1953 (BMNH).
Distribution. — Cape Province, India, Niger,
Nigeria, South Africa, Sudan, Upper Volta, and
Zaire.
Differential diagnosis. — i) frontal tubercles
small, rounded, ii) supraalar 1, iii) spurs of mid and
hind tibia unequal, iv) gonocoxite and gonostylus
short and broad and v) superior volsella digitiform
with 2 long setae at apex and a few small setae over
it.
Remarks. - The specimens collected from the
rice fields conform fully with those of Freeman
(1954), except minor morphometrics.
11. Dicrotendipes pelochloris (Kieffer)
Tendipes pelochloris Kieffer, 1912: 39; 1916: 113; Sub-
lette & Sublette 1973: 413 (listed in an unplaced series
of Chironomini).
Dicrotendipes pelochloris; Epler 1988: 134.
Limnochironomus niveicauda Kieffer, 1921a: 585.
Chironomus (Limnochironomus) niveicauda; Johannsen
1932: 528.
Dicrotendipes niveicauda; Sublette & Sublette 1973: 404;
Hashimoto et al. 1981: 13.
Dicrotendipes niveicaudus, Sasa & Hasegawa 1983: 321;
Sasa & Kikuchi 1986: 19.
Chironomus inferior Johannsen, 1932: 534.
Chironomus (Dicrotendipes) wirthi Freeman, 1961a: 692.
Kimius honsooi Ree, 1981: 218.
Xenochtronomus loripes Guha & Chaudhuri, 1981: 163.
Einfeldia loripes, Chaudhuri & Guha 1987: 27.
CHAUDHURI & CHATTOPADHYAY: Chironomids
Material examined. — 9 larvae, Burdwan, 7 September
1987; 6 larvae, 7 exuviae, 12 @ and 10 © pupae, 8 4 and
7 2 exuviae, 8 4 and 8 9, all reared; 2 À, Naihati, 16
September 1987.
Distribution. — India, Indonesia, Japan, Korea,
Pakistan, Philipines, Taiwan and Thailand.
Differential diagnosis. - Larva: i) plumose SI, ii)
pecten epipharyngis a single plate with 5 teeth, iii)
premandible with 2 unequal apical teeth, preman-
dibular brush with spines, iv) mandible without
pecten mandibularis, inner and outer margin with
indentations, v) mentum with dome-shaped me-
dian tooth, 5th and 6th lateral teeth appear to be
fused and vi) abdominal segment XI with 1 pair
short ventral tubules.
Pupa: 1) frontal apotome with conical cephalic
tubercles, ii) thoracic horn with a bunch of finely
branched filaments, iii) tergites V- VIII with median
shagreen interrupted in the middle, iv) segment
VIII with simple caudolateral spurs ending in a
point and v) G/F 0.70 in male and 0.67 in female.
Adult: i) frontal tubercles small, ii) scutellum with
6-8 setae, iii) wing smoky, R,,, bare, haltere yellow
and bare, iv) fore tibial scale blunt bearing 2 long
setae, v) tarsomeres II-V dark brown, vi) hypopy-
gium with gonostylus slightly attenuated at apex;
superior volsella well developed with setae arranged
throughout the body, inferior volsella gently bowed
ventrally and vii) genitalia with broad postgenital
plate, seminal capsules approximately equal.
12. Dicrotendipes septemmaculatus (Becker)
Chironomus septemmaculatus Becker, 1908: 77.
Dicrotendipes septemmaculatus, Cranston & Armitage
1988; Epler 1988: 42.
Dicrotendipes pictipennis Kieffer, 1913a: 23; Freeman
19556; 22:
Dicrotendipes pilosimanus Kieffer, 1914: 262; Freeman
1955a: 372; Sublette & Sublette 1973: 404; Reiss 1977:
93, 1978: 75, 1986; 159; Freeman & Cranston 1980:
190; Contreras-Lichtenberg 1986: 716; Chaudhuri &
Guha 1987: 27.
Chironomus (Dicrotendipes) pilosimanus,
1954: 19, 1957: 360, 1961b: 247, 196la: 694.
Dicrotendipes formosanus Kieffer, 1916: 115; Sublette &
Sublette 1973: 403; Hashimoto et al. 1981: 12; Sasa &
Hasegawa 1983: 320.
Dicrotendipes formosanus var. frontalis Kieffer, 1916:
116.
Dicrotendipes speciosus Kieffer, 1924a: 256; Kieffer
19252299!
Dicranotendipes speciosus, Kruseman 1949: 254 (mis-
spelling).
Stictochironomus sexnotatus Goetghebuer, 1930: 95.
Chironomus hirtitarsis Johannsen, 1932: 534; Sublette &
Sublette 1973: 402.
Dicrotendipes hirtitarsis; Guha et al. 1982: 30; Chaudhuri
& Guha 1987: 27 (Misspelling).
Polypedilum quatuordecimpunctatum
1936a: 488.
Dicrotendipes quatuordecimpunctatus, Contreras-Lich-
tenberg 1986: 710.
Freeman
Goetghebuer,
157
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Chironomus (Dicrotendipes) pilosimanus subsp. guatuor-
decimpunctatus, Freeman 1957: 361.
Dicrotendipes rajasthani Singh & Kulshrestha, 1977: 233.
Material examined. - 4 4, Burdwan, 6 September 1987;
5 6,9 © Farakka, 29 September 1986.
Distribution. — Algeria, Australia, Bangladesh,
Burma, Canary Islands, Egypt, Ethiopia, India, Is-
rael, Japan, Kenya, Lebanon, Malagasy, Malaysia,
N. Rhodesia, S. Rhodesia, Spain, Sri Lanka, Sudan,
Sumatra, Taiwan, Thailand „Transvaal, Uganda, Ye-
men and Zaire.
Differential diagnosis. — i) frontal tubercles well
developed, ii) scutellum with 12 setae in two rows,
ili) wing with 6 grey spots, iv) anal point with
broader apex, v) shape of gonostylus, vi) superior
volsella bent inward near apex bearing 3-4 apical
setae and vii) inferior velsella bifurcated into 2
blunt lobes, larger one with 6-8 and smaller one
with 5-6 setae.
13. Endochironomus pekanus (Kieffer)
Tendipes pekanus Kieffer, 1916: 105.
Endochironomus pekanus, Ree & Kim 1981: 146.
Material examined. - 2 4, Farakka, 29 September 1986:
5 6, Naihati, 16 September 1987.
Distribution. - Korea, Taiwan and India (present
record).
Differential diagnosis. — i) frontal tubercles ab-
sent, ii) fore tibial scale with a sharp pointed spur
and 3 long setae, iii) pulvilli moderately developed,
iv) characteristic dark stripes on abdominal ter-
gites, v) superior volsella dark with pallor apex and
vi) inferior volsella dark brown with 15-17
branched incurved setae.
Remarks. - The curved setae on the inferior
volsella, as reported by Ree & Kim (1981) appear
to be branched in the specimens recorded here.
Otherwise the specific characters conform with the
Korean specimens.
14. Harnischia acuta (Goetghebuer)
Chironomus (Harnischia) acutus Goetghebuer, 1936a:
470.
Cryptochironomus acutus, Freeman 1955b: 17.
Chironomus (Cryptochironomus) acutus; Freeman 1957:
STE
Harnischia acuta, Sasa & Hasegawa 1983: 323.
Material examined. - 1 8, Kalyani, 21 August 1986; 4
6, Howrah, 4 September 1987; 2 &, Burdwan, 6 Sep-
tember 1987; 1 & paratype (B.M. 1957-264). Transvaal,
Neispruit, G. H. Frank (BMNH).
Distribution. — Belgian Congo, Cape Province,
Japan, New Zealand, Nigeria and India (present
record).
Differential diagnosis. — i) eye gothic shaped
158
without a dorsal extension, ii) fore tibia with a blunt
scale bearing 3 long setae, iii) colour pattern of
tarsometers of fore, mid and hind legs, iv) anal
point slender with rounded apex, v) gonocoxite
with a setose lobe and gonostylus with swollen base
and vi) superior volsella reduced and inferior vol-
sella rudimentary.
Remarks. - A study of a paratype specimen from
BMNH, London, and comparison with collected
specimens reveal that the species should be consi-
dered as Harnischia acuta (Goetghebuer).
15. Harnischia incidata Townes
Harnischia incidata Townes, 1945: 166; Hashimoto et al.
1981: 22.
Material examined. - 5 4, Berhampur, 17 August, 1986;
3 6, Palasi, 18 August, 1986.
Distribution. - North America, Thailand and In-
dia (present record).
Differential diagnosis. - i) frontal tubercles well
marked, ii) antepronotum reduced, iii) R,+; with 1
apical seta, iv) haltere light yellow with 6-8 setae,
v) spurs of mid tibia unequal and spurs of hind tibia
equal, vi) pulvilli small, vii) well marked gonocoxite
and gonostylus and viii) superior volsella reduced
and inferior volsella absent.
Remarks. - The frontal tubercles recorded in
Indian specimens were probably overlooked in the
specimens from Thailand (Hashimoto et al. 1981).
16. Harnischia tenuitubercula sp. n.
Type material: Holotype g, India, West Bengal, Fa-
rakka, 29 September 1986, leg. S. Chattopadhyay (Type
no. 183, B.U. Ent.). Paratypes: 3 @, data as holotype.
Differential diagnosis. — i) antepronotum well
developed with emargination in the middle, ii)
prescutellars present, iii) ratio of spurs of mid and
hind tibia equal, iv) tongue shaped anal point, v)
gonostylus oblong with abruptly pointed apex and
vi) superior volsella rudimentary.
Remarks. - The species is named tenwitubercula
because of its small frontal tubercle. It appears to be
close to H. acuta (Goetghebuer) in body coloura-
tion. In its gonocoxite and gonostylus, this species
also resembles H. argentea Townes, 1945.
Description
Adult. - Male: Body 2.27 (2.20-2.35, n = 4) long,
wing 1.14 (1.13-1.14, n = 6) long and 0.36 (0.35-
0.38, n = 6) wide.
Head: Brown. Vertex with 7-8 setae (IV 2-3, OV
2, PO 3). Corona bare. Clypeus with 10-12 setae,
clypeal ratio 0.86. Maxillary palp pale brown, ratio
of palpomere length I-V: 5 : 6 : 27 : 22: 23, L/W
3.8. Eyes gothic shaped with a dorsal extension of
0.018 long. Frontal tubercles (fig. 22) small. An-
tenna brown, ratio of flagellomere length I-XI :
1228260362 98292929:2929 3 Gor AVR Does
pedicel ratio 1.09. CA 0.52; CP 1.40.
Thorax: Yellow. Antepronotum well developed
with emargination in the middle, antepronotals 0.
Mesonotum with 3 brown vittae. Acrostichals 10-
12 biserial, dorsocentrals 7-8, prealars 3, prescutel-
lars 2. Scutellum with 8 setae in a transverse row,
postscutellum dark brown and bare.
Wing (fig. 23): Hyaline. Brachiolum with 2 setae
and 8-9 sensilla campaniformia. R with 19-20, R,
with 12-13 and R‚‚; with 17-18 setae. R,;, meets
C at a distance of 0.028 from R,; RM faint and
proximal to FCu. An ends proximal to FCu. Squama
with 4 setae. Haltere light yellow and bare. CR 0.99;
WAR TIA
Legs: Yellow. Fore tibia with a blunt scale (fig.
24) bearing 2 long setae. Spurs of mid tibia unequal,
0.027 and 0.018 long, ratio of length of spurs to the
apical diameter of mid tibia 9 : 11 and 6 : 11; spurs
of hind tibia unequal, 0.027 and 0.018 long, ratio of
length of spurs to the apical diameter of hind tibia
9 : 13 and6 : 13. Sensilla chaetica indistinguishable.
Pulvilli moderately developed. Proportions and ra-
uos of leg-segments in table 1.
Abdomen: Light yellow. Hypopygium (fig. 25)
with tongue-shaped setose anal point 0.042 long
and distinctly marked gonocoxite and gonostylus.
Gonocoxite with 11 setae; gonostylus oblong with
abruptly pointed apex bearing 9 setae at its inner
margin and 13-14 setae over it. Superior volsella
rudimentary bearing 2 small setae, inferior volsella
absent. Transverse sternapodeme 0.06, lateral
sternapodeme 0.09, coxapodeme 0.039 and phallap-
odeme 0.06 long. HR 0.69; HV 1.6.
Female. - Unknown.
Distribution. - India.
17. Harnischia viridula (Linnaeus)
Tipula viridula Linnaeus, 1767: 975.
Chironomus (Harnischia) viridulus, Goetghebuer 1928:
86.
Chironomus (Chironomus) viridulus, Edwards 1929: 390.
Cryptocladopelma viridula; Pinder 1978: 18.
Harnischia viridula, Hashimoto et al. 1981: 21; Sasa &
Hasegawa 1983: 324; Sasa 1985: 35; Sasa & Kikuchi
1986: 20.
Material examined. - 5 &, Burdwan 30 May 1986; 3 4,
Chinsura, 5 February 1987.
Distribution. - England, Japan, North America,
Sweden, Thailand and India (present record).
Differential diagnosis. — i) frontal tubercles well
developed, ii) postscutellum bearing 2 setae, iii)
R,4; with 1-2 apical setae, iv) pulvilli well deve-
CHAUDHURI & CHATTOPADHYAY: Chironomids
loped, v) unmarked gonocoxite and gonostylus and
vi) superior volsella rudimentary bearing 2 setae.
18. Kiefferulus barbatitarsis (Kieffer)
Chironomus barbatitarsis Kieffer, 1911: 154.
Kiefferulus barbatitarsis; Chaudhuri & Ghosh 1986: 277;
Chaudhuri & Guha 1987: 28.
Material examined. — 6 larvae, Uttarpara, 9 September
1987; 7 larvae, 8 & and 6 9 pupae, 4 @ and 5 9 exuviae,
6 8 and 5 9 all reared; 5 @, Howrah, 4 September 1987;
4 8, Kalyani, 10 February 1986; 9 &, Hooghly, 4 Sep-
tember 1987.
Distribution. — India.
Differential diagnosis. — Larva: i) S I of labrum
densely pectinate, ii) pecten epipharyngis with 18
unequal teeth, iii) ventromental plate with
wrinkled outer surface and iv) posterior parapods
with 15-16 well-sclerotized claws.
Pupa: 1) thoracic horn with a bunch of fine fila-
ments, and ii) segment VIII with 1 pair of caudo-
lateral spurs, each with 4-5 teeth in male and 8-10
in female.
Adult: 1) scutellum with 20 setae in two trans-
verse rows, ii) branchiolum with 4 setae and iii)
hypopygium with tongue-like anal point, superior
volsella bent with subacute apex and bare and infe-
rior volsella curved with blunt apex bearing numer-
ous curved setae.
19. Kiefferulus calligaster (Kieffer)
Chironomus calligaster Kieffer, 1911: 160.
Kiefferulus calligaster; Chaudhuri & Ghosh 1986: 285;
Chaudhuri & Guha 1987: 28.
Material examined. — 5 larvae, Burdwan, 14 September
1986; 6 larvae, 6 & and 7 Q pupae, 5 and 6 Q exuviae,
9 8 and 6 9 all reared; 2 4, Kalyani, 26 October 1986; 5
8, Kalna, 4 February 1988.
Distribution. — India.
Differential diagnosis. - Larva: i) S I of labrum
plumose, ii) premandible with 5 curved apical teeth
and well developed premandibular brush, iii) ven-
tromental plate with smooth outer surface, and iv)
posterior parapods long, each with 13-14 claws.
Pupa: 1) frontal apotome with 1 pair of funnel-
shaped cephalic tubercles, ii) thoracic horn with 2
plumose branches, 111) tergite VII without shagreen
and iv) segment VIII with 1 pair blunt, weak caudo-
lateral spurs.
Adult: i) scutellum with 6 setae in a single trans-
verse row, ii) brachiolum with 2 setae, iii) fore
femur with apical band, iv) pulvilli large and broad
and v) hypopygium with dark peglike anal point,
and superior volsella with swollen base bearing 5
setae.
159
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
20. Microchironomus fuscitarsus (Guha &
Chaudhuri)
Cryptochironomus fuscitarsus Guha & Chaudhuri, 1979:
Dy.
Microchironomus fuscitarsus, Chaudhuri & Chattopad-
hyay 1988: 175.
Material examined. - 7 larvae, Howrah, 8 March 1987;
3 larvae, 8 6 and 8 © pupae, 5 @ and 7 9, exuviae, 8 G and
8 9, all reared; 3 6, Burdwan, 6 September 1987.
Distribution. - India.
Differential diagnosis. — Larva: i) blade-like S I
and SII, ii) premandible with 2 unequal apical teeth,
iii) mandible with well developed pecten mandibu-
laris and iv) fan-shaped ventromental plate with
wrinkled outer margin.
Pupa: 1) conical cephalic tubercles without fron-
tal seta, ii) thoracic horn with 4 main bunch of
finely branched filaments, iii) tergites I, II and VII
bare and iv) caudolateral spur on segment VIII
simple with sharp point.
Adult: 1) antepronotum with 5 antepronotals, ii)
scutellum with 10 setae, iii) fore tibial scale with 2
long setae and iv) hypopygium with small setaceous
lobe basolateral to anal point; superior volsella with
1 apical seta.
21. Microchironomus tener (Kieffer)
Chironomus tener Kieffer, 1918: 48.
Tendipes (Parachironomus) tener; Kruseman 1933: 125,
190-191.
Leptochironomus tener; Kugler 1971: 341; Shilova 1976:
118-119; Pinder 1978: 126.
Microchironomus tener; Saether 1977: 101; Freeman &
Cranston 1980: 193; Moller-Pillot 1984: 244; Chaud-
huri & Chattopadhyay 1988: 177.
Parachironomus tener, Hashimoto et al. 1981: 19.
Cryptochironomus aegyptus Kieffer, 1925: 288.
Chironomus (Cryptochironomus) balticus Pagast, 1931:
218.
Chironomus forcipatus Freeman, 1954: 20; 1957: 394;
1961a: 699.
Material examined. — 9 larvae, Kalyani, 17 February
1987; 5 larvae, 6 @ and 5 Q pupae, 4 & and 5 Q exuviae,
6 è and 3 9, all reared.
Distribution. — Australia, Chad, Ghana, India,
Madagascar, Poland, Rhodesia, South Africa and
Zaire.
Differential diagnosis. - Larva: i) pecten epipha-
ryngis indistinct, ii) mandible with 3 flat lateral
teeth; seta interna with 3-4 filaments; pecten man-
dibularis absent, iii) mentum with minute 4th lat-
eral tooth and iv) ventromental plate fan-shaped
with faint rays ending before the margin.
Pupa: 1) elevated cephalic tubercle with an apical
seta, 11) tergites I and II bare, and iii) pattern of
shagreen on tergites and lateral setae on segments.
Adult: i) frontal tubercles absent, ii) haltere with
160
7-10 setae and iti) hypopygium with stout anal
point and large setaceous lobe basolaterally; super-
ior volsella long and slender with 2-3 setae at apex.
22. Paracladopelma aratra sp. n.
Type material. - Holotype @, India, West Bengal, Bally,
9 March 1987, leg. S. Chattopadhyay (Type no. 184, B.U.
Ent.). Paratypes: 4 @, data as holotype.
Differential diagnosis. — 1) frontal tubercles ab-
sent; ii) prescutellar 1, iii) R,,; with 2 setae at the
apex, iv) spurs of hind tibia equal, v) superior vol-
sella strongly sclerotised appearing as an outgrowth
of the gonostylus, inferior volsella plough-like and
vi) anal point tubular.
Remarks. - Paracladopelma aratra is named ara-
tra because of its plough-shaped inferior volsella of
the male hypopygium. In the structure of the gono-
coxite and gonostylus of the male hypopygium, the
present species resembles P. graminicolor
(Kieffer). The anal point of the male hypopygium
shows similarity to that of P. brincki (Freeman).
Description
Adult. - Male: Body 2.09 (2.06-2.11, n= 5) long,
wing 0.91 (0.91-0.92, n = 6) long and 0.31 (0.29-
0.31, n = 6) wide.
Head: Brown. Vertex with 7-9 setae (IV 2-3, OV
3-4, PO 2). Corona bare. Clypeus with 10-12 setae,
clypeal ratio 0.94. Maxillary palp pale brown, ratio
of palpomere length I-V: 9 : 6 : 19 : 24 : 33; L/W
3.8. Eyes reniform without a dorsal extension.
Frontal tubercles absent. Antenna brown, ratio of
flasellomerelene th EX EM 2 ED EICECEEE
7 :7 :7 : 102; AR 1.5; pedicel ratio 1.2 CA 0.66; CP
1.23
Thorax: Yellow. Antepronotum well developed
without emargination, antepronotals O. Mesono-
tum with 3 brown vittae. Acrostichals 4-6 irregu-
larly biserial, dorsocentrals 5-6 uniserial, prescutel-
lar 1, prealars 3. Scutellum with 4-6 setae,
postscutellum dark brown and bare.
Wing (fig. 26): Opaque, veins light brown. Bra-
chiolum with 1 seta and 16-18 sensilla campanifor-
mia. R and R, without seta; R,;; with 2 setae at the
apex; R,;, meets C at distance of 0.981 from R,;
RM faint and proximal to FCu. Squama with 5-6
setae. Haltere light yellow with 9-11 setae. CR 0.95;
WI 125,
Legs: Yellow. Fore tibia with a blunt scale (fig. 27)
bearing 2 long setae. Spurs of mid tibia equal 0.018
long, ratio of length of spurs to the apical diameter
of mid tibia 6 :9; spurs of hind tibia equal 0.021 long,
ratio of length of spurs to the apical diameter of hind
tibia 7 : 11. Femur, tarsomere 1 of fore leg and
tarsomere V of mid and hind leg brown. Sensilla
chactica indistinguishable. Pulvilli well developed.
Proportions and ratios of leg-segments in table 1.
Abdomen: Yellow. Hypopygium (fig. 28) with
tubular anal point 0.036 long and with 3-5 basolat-
eral setae. Gonocoxite with 6-7 gonostylus short
and stout bearing 8 setae at its.inner margin and 1 1-
12 setae over it. Superior volsella strongly sclero-
tised appearing as an outgrowth of the gonostylus;
inferior volsella more or less plough like having 2
long setae. Transverse sternapodeme 0.033, lateral
sternapodeme 0.069, coxapodeme 0.033 and phal-
lapodeme 0.036 long. HR 0.79; HV 2.71.
Female. - Unknown.
Distribution. - India.
23. Paracladopelma sacculifera sp. n.
Type material. - Holotype @, India, West Bengal, Ka-
lyani, 17 February, leg. S. Chattopadhyay (Type no. 185,
B.U. Ent.). Paratypes: 5 @, data as holotype.
Differential diagnosis. - i) frontal tubercles very
small, it) antepronotum very thin without dorsal
emergination, ii) spur of hind tibia unequal, iv)
superior volsella short and broad, inferior volsella
finger like and each with 1 long seta and v) anal
point with saccular tip.
Remarks. - The name of the species is derived
from the saccular tip of the anal point of the male
hypopygium. The species seems to be nearer to P.
camptobolis (Kieffer), P. nats (Townes) and P. doris
(Townes) in respect of the anal point, but to P.
undine (Townes) in view of the gonostylus of the
male hypopygium.
Description
Adult. - Male: Body 2.77 (2.79-2.88, n = 6) long,
wing 1.58 (1.58-1-60, n = 6) long and 0.49 (0.48-
0.49, n = 6) wide.
Head: Brown. Vertex with 12-13 setae (IV 3-4,
OV 5-6, PO 4). Clypeus with 18-19 setae, clypeal
ratio 0.92. Maxillary palp pale brown, ratio of pal-
pomeretlens the VEMOE 164652 2555 07/1
4.86. Eyes reniform without a dorsal extension.
Frontal tubercles (fig. 29) very small. Antenna
brown, ratio of flagellomere length I-XI : 12 : 7:
CHOO) ARE903 pedicel
ratio 1.09. CA 0.53; CP 0.92.
Thorax: Yellow. Antepronotum very thin with-
out dorsal emargination, antepronotals O. Meso-
notum with 3 brown vittae. Acrostichals 10-11 ir-
regularly biserial, dorsocentrals 7-8 uniserial,
prealars 4. Scutellum with 10 setae, postscutellum
dark brown and bare.
Wing (fig. 30): Opaque, veins light brown. Bra-
chiolum with 1 seta 10-12 sensilla campaniformia.
R with 7-9 setae, R, and R,,; without seta. R,,,
meets C at a distance of 0.05 from R,; RM faint and
proximal to FCu; An ends below FCu. Squama with
CHAUDHURI & CHATTOPADHYAY: Chironomids
5-6 setae. Haltere yellowish green with 6-7 setae.
CR°0192: VR 1.19.
Legs: Yellow. Fore tibia with a blunt scale (fig.
31) bearing 2 long setae. Spurs of mid tibia subequal
0.018 and 0.015 long, ratio of length of spurs to the
apical diameter of mid tibia 6 : 13 and 5 : 13; spurs
of hind tibia unequal 0.021 and 0.012 long, ratio of
length of spurs to the apical diameter of hind tibia
7:15 and 4 : 15. Apex of fore tibia and tarsomeres
of fore, mid and hind legs brown. Sensilla chaetica
indistinguishable. Pulvilli well developed. Propor-
tions and ratios of leg-segments in table 1.
Abdomen: Yellow. Hypopygium (fig. 32) with
anal point with saccular apex bearing 8-9 setae at
its base. Gonocoxite short with 8-10 setae; gono-
stylus long and little incurved with 11-13 small
setae in its inner margin and 15-17 setae over it.
Superior volsella short, broad and pubescent bear-
ing | long seta; inferior volsella finger like bearing
1 long and numerous minute setae. Transverse ster-
napodeme 0.042, lateral sternapodeme 0.087, cox-
apodeme 0.054 and phallapodeme 0.042 long. HR
0.65; HV 1.98.
Female. - Unknown.
Distribution. - India.
24. Pentapedilum robusticeps Guha &
Chaudhuri
Pentapedilum robusticeps Guha & Chaudhuri, 1985: 195.
Material examined. - 4 4, Palasi, 18 August 1986; 5 &,
Uttarpara, 24 February 1987.
Distribution. — India.
Differential diagnosis. — i) scutellum with 26
setae in two transverse rows, ii) brachiolum with 7
setae and iii) hypopygium with robust anal point,
stout gonostylus, narrow superior volsella and fin-
ger like inferior volsella.
25. Pentapedilum uncinatum (Goetghebuer)
Tanytarsus uncinatus Goetghebuer, 1921: 110.
Pentapedilum uncinatum, Edwards 1929: 376; Pinder
1978: 134; Sasa & Kikuchi 1986: 21.
Material examined: 7 8, Kalyani, 17 February 1987; 1
6, Spain, Levida, 6 July 1973, Coll. P. S. Cranston
(BMNH).
Distribution. - Belgium, England, Germany, Ja-
pan, Spain and India (present record).
Differential diagnosis. — i) frontal tubercles ab-
sent, ii) fore tibia with a blunt scale bearing 3 setae,
ii) length of mid and hind tibia equal, iv) pulvilli
moderately developed, v) superior volsella with 1
long lateral seta arising at about the distal 2/3 from
the base.
161
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
26. Polypedilum aegyptium Kieffer
Polypedilum aegyptium Kieffer, 1925: 270; Freeman
1958: 281; Guha & Chaudhuri 1983: 635; Chaudhuri &
Guha 1987: 28.
Polypedilum iris Goetghebuer, 1937: 61.
Polypedilum airense Freeman, 1956: 96.
Material examined. - 5 8, Chinsura, 6 February 1987.
Distribution. - Africa, Egypt and India.
Differential diagnosis. - i) frontal tubercles ab-
sent, ii) thorax dark brown, iii) triangular fore tibial
scale having a sharp spine, iv) wing with brown
markings and clouds and v) superior volsella more
angular.
27. Polypedilum annulatipes (Kieffer)
Chironomus annulatipes Kieffer, 1910: 234.
Polypedilum annulatipes, Kieffer 1921b: 100, 1922: 30;
Freeman 1958: 277-278; Sublette & Sublette 1973: 407;
Chaudhuri et al. 1981: 115-117; Chaudhuri & Guha
1987: 28.
Polypedilum octostictum Kieffer, 1921b: 101, 1922: 39.
Polypedilum octomaculatum Goetghebuer, 1934: 195,
1936a: 488.
Material examined. - 6 @, Bally, 13 February 1986; 1 6,
India, Orissa, Barkuda Lake, 1 October 1922, leg. N. An-
nadale (NZC, Calcutta).
Distribution. - Africa, Egypt, India, Pakistan and
Zaire.
Differential diagnosis. — i) presence of single
spur in mid and hind tibia, ii) absence of frontal
tubercles, iii) antepronotum not collar-like and iv)
abdominal segment VIII constricted at the base.
Remarks. - From the present study it may be
inferred that P. annulatipes is a distinct species in
the genus Polypedilum Kieffer.
28. Polypedilum ascium Chaudhuri, Guha & Das
Gupta
Polypedilum ascium Chaudhuri, Guha & Das Gupta, 1981:
117-119; Chaudhuri & Guha 1987: 28.
Material examined. - 4 @, Palasi, 18 August 1986; 1 8,
Chinsura, 6 February 1987.
Distribution. - Bhutan and India.
Differential diagnosis. — i) mesonotum brown
with dark brown patches, ii) antepronotum very
thin, iii) scutellum with 6 setae, iv) fore tibial scale
elongated and blunt and v) inferior volsella axe-
shaped.
29. Polypedilum chaudhurii Chaudhuri, Guha &
Das Gupta
Polypedilum chaudhuri Chaudhuri, Guha & Das Gupta,
1981: 119-122; Chaudhuri & Guha 1987: 28.
162
Material examined. — 16 8, Naihati, 16 September
1987.
Distribution. — India.
Differential diagnosis. — i) colour pattern of
wing, ii) colour bands of the legs, iii) two rows of
scutellar setae, iv) spur of mid tibia bent and v)
hypopygium with anal point bearing two projec-
tions, superior volsella with uniform width, gently
bent at the middle bearing 4 setae and numerous
weak setae and elongated, little bowed inferior vol-
sella.
30. Polypedilum circulum sp. n.
Type material. - Holotype 8, India, West Bengal, Ka-
lyani, 25 October 1986, leg. S. Chattopadhyay (Type no.
186, B.U. Ent.). Paratypes: 8 8, data as holotype.
Differential diagnosis. — i) frontal tubercles ab-
sent, ii) antepronotum transparent with some cone
shaped apex and emargination in the middle, iii)
haltere dark brown with 6-8 setae, iv) spurs of mid
and hind tibia equal, v) colour pattern of the abdo-
men, vi) anal point slender, ending just before tip
of appendage 2, vii) gonocoxite short, gonostylus
clavate with 6 long setae on its inner margin, viii)
superior volsella curved with blunt tip and 1 long
lateral seta near the base.
Remarks. - Polypedilum circulum is named after
the slender appendage 1 of the male hypopygium.
The present species resembles P. nubeculosum
(Meigen) and P. youngosanensis Ree & Kim in the
gonostylus and P. angustum Townes in appendage
1. The anal point of the male hypopygium also
shows similarities with that of P. acutum Kieffer
and P. angustum Townes.
Description
Adult. - Male: Body 3.37 (2.29-3.58, n= 9) long,
wing 1.76 (1.75-1.79, n = 10) long and 0.53 (0.52-
0.55, n = 10) wide.
Head: Brown. Vertex with 15-17 setae (IV 4-5,
OV 7-8, PO 4). Corona bare. Clypeus with 17-19
setae, clypeal ratio 0.85. Maxillary palp pale brown,
ratio of palpomere length I-V: 10 : 8 : 19 : 24 : 35;
L/W 3.80. Eyes reniform with a dorsal extension of
0.01 long. Frontal tubercles absent. Antenna brown,
ratio of flagellomere length I-XII: 19: 7:8:7:
Ties OFM] Ha eNO AINE ZA pedicel
ratio 0.85. CA 0.50; CP 1.55.
Thorax: Brown. Antepronotum transparent
with cone shaped apex and slight emergination in
the middle, antepronotals O. Mesonotum brown.
Acrostichals 10-11 irregularly biserial, dorsocen-
trals 8-9 uniserial, prealars 2-3. Scutellum with 6
setae in a transverse row, postscutellum dark brown
and bare.
Wing (fig. 33): Hyaline. Brachiolum with 1 seta
and 21-23 sensilla campaniformia. R with 17-18, R,
with 11-12 and R,;; with 19 setae; R,,; meets Cat
a distance of 0.04 from R,; RM pale and proximal
to FCu. An ends proximal to FCu. Squama with 8
setae. Haltere dark brown with 6-8 setae. CR 0.98;
VR 1.16.
Legs: Yelllow. Fore tibia with a triangular scale
(fig. 54) bearing a very short apical spine and 2 long
setae. Mid tibia with single spur 0.027 long, ratio of
length of spur to the apical diameter of mid tibia 9 :
13, hind tibia with single spur 0.027 long, ratio of
length of spur to the apical diameter of hind tibia
9 : 15. Basal and apical portion of fore tibia dark
brown, tarsomeres uniform in colour; tarsomere V
of mid and hind leg yellow to pale brown. Sensilla
chaetica indistinguishable. Pulvilli moderately
developed. Proportions and ratios of leg-segments
in table 1.
Abdomen (fig. 35): Tergites yellowish brown
with dark brown band along lateral margin, tergites
UI and VI with dark brown median patches in
addition to the above. Hypopygium (fig. 36) with
tubular anal point 0.042 long, subequal to appen-
dage 2 and with 4-5 basolateral setae. Gonocoxite
short, stout with 7-9 setae; gonostylus clavate with
1 apical seta, 6 long setae on its inner margin and
12-14 setae over it. Superior volsella slender, bent
inward with 3 basal setae and 1 long lateral seta
near the base; inferior volsella stout, linear with 11-
12 incurved setae besides single long apical seta.
Transverse sternapodeme 0.036, lateral sternapo-
deme 0.066, coxapodeme 0.048 and phallapodeme
0.075 long. HR 0.44; HV 1.27.
31. Polypedilum lineatum Chaudhuri, Guha &
Das Gupta
Polypedilum lineatum Chaudhuri, Guha & Das Gupta,
1981: 129-131; Chaudhuri & Guha 1987: 28.
Material examined. - 9 &, Haldia, 16 October 1987.
Distribution. - India.
Differential diagnosis. i) corona with setae, ii)
frontal tubercles absent, iii) antepronotum dark,
very thin without dorsal emergination, and super-
ior volsella well developed, wider at the apex, with
numerous small setae.
52. Polypedilum nubifer (Skuse)
Chironomus nubifer Skuse, 1889: 249; Kieffer 1906: 20;
1917: 205.
Polypedilum (Polypedilum) nubifer, Freeman 1961a; 707.
Polypedilum nubifer, Martin 1966: 157; Sublette & Sub-
lette 1973: 408; Porter & Martin 1977: 41; Sasa &
Sublette 1980: 33; Hashimoto et al. 1981: 23; Ree &
Kim 1981: 158; Sasa & Hasegawa 1983: 327; Sasa &
Kikuchi 1986: 26; Chaudhuri & Guha 1987: 28; Chat-
topadhyay et al. 1988: 29.
CHAUDHURI & CHATTOPADHYAY: Chironomids
Chironomus triparticus Kieffer, 1910: 231 (see Freeman
de ceylonicus Kieffer, 1911: 136 (see Freeman
DE pelostolum Kieffer, 1912: 41 (see Freeman
ae pharao Kieffer, 1925: 274 (see Freeman
(Polypedilum) albiceps Johannsen, 1932:
Di (Polypedilum) albiceps, Tokunaga 1964:
rate ae (Polypedilum) octoguttatus Tokunaga, 1936:
ee octoguttatum; Sasa 1979: 15.
Material examined. — 5 larvae, Burdwan, 14 September
1986; 8 larvae, 4 @ and 5 9 pupae, 6 8 and 5 2 exuviae,
7 & and 7 9, all reared; 2 6, Kalyani, 10 February 1986;
3 À, Chinsura, 6 February 1987; 5 6, Hooghly, 4 Sep-
tember 1987; 2 8, Uttarpara, 9 September 1987.
Distribution. - Africa, Australia, Egypt, Indone-
sia, India, Iraq, Japan, Korea, Micronesia, Morocco,
Sri Lanka and Taiwan.
Differential diagnosis. - Larva: S I and S II plu-
mose, ii) pecten epipharyngis comprising 3 toothed
plates, ili) premandible with 2 unequal apical teeth,
iv) seta interna with 3 main plumose branches and
v) mentum with short Ist and 7th lateral teeth.
Pupa: 1) frontal apotome with elevated cephalic
tubercles, ii) thoracic horn with 4 main bunches of
finely branched filaments, iii) tergites I and VIII
bare, iv) absence of dorsal seta on segment VIII and
v) caudolateral spur of segment VIII with 14-24
variable, stout spines.
Adult: i) frontal tubercles small, ii) wing with 10
spots, iii) squama fringed, and iv) hypopygium with
slender anal point, gonostylus attenuated with 1
long seta at apex; inferior volsella with 1 long seta
at apex.
33. Polypedilum medivittatum Tokunaga
Polypedilum (Polypedilum) medivittatum Tokunaga,
1964: 588; Sasa & Hasegawa 1983: 329.
Chironomus (Polypedilum) sp. no. 17, Johannsen 1932:
524.
Material examined. - 6 8, Kalyani, 26 October 1986.
Distribution. - Caroline Islands (Palau, Yap and
Panope), Japan, S. Mariana Islands (Gham), South
Sumatra and India (present record.)
Differential diagnosis. - i) antepronotum narrow
without emargination, ii) scutellum with 2-4 setae,
iii) haltere yellow with 5 setae, iv) pulvilli well
developed, v) tergites I-VI light yellow and tergite
VIII brown, vi) superior volsella scimitar like and
vii) inferior volsella not beyond tip of anal point.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
34. Polypedilum suturalis (Johannsen)
Chironomus (Polypedilum) suturalis Johannsen, 1932:
522:
Polypedilum suturalis, Sublette & Sublette 1973: 408; Ha-
shimoto et al. 1981: 25.
Material examined. - 6 g, Haldia, 16 October 1987.
Distribution. - South Sumatra, Thailand and In-
dia (present record).
Differential diagnosis. - i) prescutellar 1, ii) fore
tibial scale triangular bearing a short subapical
spine and 2 long setae, iii) apical diameter of mid
and hind tibiae is lesser than the length of spurs, iv)
colour pattern of abdomen, v) anal point long with
pointed inwardly bent apex, vi) superior volsella
very slender, arcuate with 1 seta on subbasal part
and 11-12 minute setae and vii) inferior volsella not
reaching tip of the anal point.
35. Polypedilum yapensis Tokunaga
Polypedilum (Polypedilum) yapensis Tokunaga, 1964:
595; Hashimoto et al. 1981: 25.
Material examined. - 9 4, Howrah, 8 March 1987; 3 4,
Bally, 8 March 1987.
Distribution. — Caroline Islands (Palau, Yap),
Thailand and India (present record).
Differential diagnosis. — i) frontal tubercles well
developed, setose, ii) scutellum with 4 setae, iii) fore
tibia with a blunt scale bearing 2 setae, iv) length
of spur of mid and hind tibia equal, v) superior
volsella slender and with 1 long seta behind tip and
vi) anal point slender ending before the tip of in-
ferior volsella.
Remarks. - The Indian specimens are slightly
different from the Micronesian specimens in body
colouration, but the main specific characters seem
to be identical. The frontal tubercle, which has been
reported to be absent by Hashimoto et al. (1981),
has been noted in the present specimens.
36. Stenochironomus hilaris (Walker)
Chironomus hilaris Walker, 1848: 17.
Stenochironomus hilaris, Townes 1945: 90; Beck & Beck
1970: 33; Gillespie 1974: 238.
Stenochironomus (Stenochironomus) hilaris, Borkent
1984: 69.
Chironomus taeniapennis Coquillett, 1901: 607; Jo-
hannsen 1905: 203; Malloch 1915: 430.
Chironomus (Stenochironomus) taeniapennis; Johannsen
1957.28:
Stenochironomus taeniapennis, Townes 1945: 90 (in
part).
Chironomus nephopterus Mitchell, 1908: 7; Johannsen
1908: 280; Malloch 1915: 429.
Chironomus nephoterus [Sic]; Johannsen 1908: 280.
Stenochironomus nephopterus, Townes 1945: 90.
Chironomus zonopterus Mitchell, 1908: 12.
Stenochironomus zonopterus, Townes 1945: 91.
164
Chironomus exquisitus Mitchell, 1908: 11.
Chironomus (Stenochironomus) equisitus, Johannsen
1937: 28.
Stenochironomus exquisitus, Townes 1945: 91.
Material examined. - 4 4, Farakka, 29 September 1986.
Distribution. - Canada, U.S.A. and India (present
record).
Differential diagnosis. - i) high value of L/W
(7.14), ii) scutellum with 30 setae in two transverse
rows, ili) wing with lightly pigmented medial and
apical bands, iv) colour pattern of legs, v) fore tibia
with a blunt scale bearing 4 long setae, vi) dark anal
point with hyaline expanded tip and vii) superior
volsella short with 6 setae.
37. Stenochironomus longipalpis (Kieffer)
Tendipes longipalpis Kieffer, 1913b: 132.
Stenochironomus longipalpis; Sublette & Sublette 1973:
409.
Material examined. - 3 @, Palasi, 18 August 1986.
Differential diagnosis. — i) scutellum with 14
setae in a transverse row, ii) wing unmarked, iii)
fore tibia with an elongated scale bearing a short
spur and 4 long setae, iv) superior volsella short and
inconspicuous bearing 5-6 setae and v) inferior vol-
sella with 3 long curved setae on inner apical mar-
gin.
Remarks. - As the type is reported to be lost from
NZC, Calcutta, the species is here redescribed in
order to fix its identity.
Redescription
Adult. - Male: Body 3.51 (3.43-3.68, n = 3) long,
wing 1.48 (1.45-1.50, n = 6) long and 0.38 (0.37-
0.39, n = 6) wide.
Head: Brown. Vertex with 12-17 setae (IV 4-6,
OV 8-10, PO 0-1). Corona bare. Clypeus with 18-20
setae, clypeal ratio 1.11. Maxillary palp brown, ratio
of palpomere length I-V: 10 : 8 : 17 : 40 : 41; L/W
2.42. Eyes reniform without a dorsal extension.
Frontal tubercles absent. Antenna brown, ratio of
palpomere length EXIGE 79
SONS DS IS NAS DRS ARMEGS pedicel
ratio 1.18. CA 0.66; CP 2.16.
Thorax: Light brown. Antepronotum reduced,
antepronotals O. Mesonotum with 4 dark vittae (1
pair dorsomesad, parallel to each other and one
each dorsolaterad). Acrostichals 16 irregularly bise-
rial, dorsocentrals 17 uniserial, humeral 0.1, prea-
lars 5-6. Scutellum with 14 setae in a transverse
row, postscutellum dark brown and bare.
Wing (fig. 37): Unmarked with pale brown
veins. Brachiolum with 2 setae and 14-15 sensilla
companiformia. R with 30-34, R, with 31-35 and
R,+; with 64-70 setae; R,,; ending close to R,; RM
brown, thick and proximal to FCu. An ends below
FCu. Squama with 7-8 setae. Haltere yellow with 6-
Insetaene RI SVRSINT:
Legs: Yellow. Fore tibia with an elongated scale
(fig. 38), armed with a short spur, 0.003 long bear-
ing 4 long setae. Spurs of mid tibia subequal 0.027
and 0.024 long, ratio of length of spurs to the apical
diameter of hind tibia 9 : 15 and 8 : 15; spurs of
hind tibia also subequal 0.024 and 0.021 long, ratio
of length of spurs to the apical diameter and hind
tibia 8 : 16 and 7 : 16. Apex of fore femur brown,
fore tibia dark at base and apex but mid and hind
tibiae brown only at base. Tarsomeres broken in all
specimens.
Abdomen (fig. 39): Uniformly setaceous with a
large number of long setae at each lateral margin.
Tergites yellow with brownish tinge except the base
of tergites I-IV with dark brown. Hypopygium (fig.
40) with dark, stout anal point, apex broad, rounded
bearing 7-8 basolateral setae. Gonocoxite with 7-9
long setae; gonostylus long, slender, evenly curved
bearing 1 long apical seta, 3-4 setae on inner margin
and 11-13 setae over it. Superior volsella short and
inconspicuous bearing 5-6 setae; inferior volsella
long, narrow and curved with an articulated spine
of 0.015 long at tip and 3 long setae on inner apical
margin. Transverse sternapodeme 0.024, lateral
sternapodeme 0.06, coxapodeme 0.03, and phall-
apodeme 0.063, long. HR 0.70, HV 2.09.
Female. - Unknown.
38. Stictochironomus affinis (Johannsen)
Chironomus (Stictochironomus) affinis Johannsen, 1932:
925:
Stictochironomus affinis, Ali et al. 1987; Chaudhuri &
Guha 1987: 29.
Materail examined. — 8 larvae, Katwa, 8 February 1987;
5 larvae, 9 G and 8 © pupae, 5 d and 6 Q exuviae, 8 8 and
4 Q, all reared; 6 6, Uttarpara, 24 February 1987; 3 8,
Berhampur, 2 January 1988.
Distribution. — India and Indonesia.
Differential diagnosis. — Larva: i) SI and S II
plumose and ii) pecten epipharyngis comprising 3
toothed plates, median with 3-5 and each lateral
with 5 teeth.
Pupa: i) tergite I with uniformly distributed spin-
ules, ii) segments II-VII with 2, 3, 3, 3, 3, 4, 4 pairs
of lateral setae and iii) caudolateral spur on segment
VII with 7-8 spines.
Adult: i) corona bare, ii) frontal tubercles pres-
ent, 111) scutellum with dark posterior border bear-
ing 24 setae, iv) haltere bare and v) hypopygium
with curved, pointed superior volsella and slender
anal point with subacute apex.
Remarks. - This species was described by Jo-
hannsen (1932) from a female, collected on Bali,
Indonesia. Ali et al. (1987) confirmed the identity
CHAUDHURI & CHATTOPADHYAY: Chironomids
of this species by comparing reared specimens with
the description by Johannsen and by examining the
type, received from BMNH, London. They also
made a detailed description of the life stages, with
a brief account of the general behaviour of larvae
and the ovipository behaviour of the female.
39. Stictochironomus obscurus (Guha &
Chaudhuri) comb. n.
Polypedilum obscurum Guha & Chaudhuri, 1983: 637.
Material examined. — 6 larvae, Burdwan 30 May 1986;
6 larvae, 8 6 and 8 © pupae, 5 @ and 5 Q exuviae, 10 9,
all reared; 2 4, Bally, 9 March 1987.
Distribution. — India.
Differential diagnosis. - Larva: i) AR 0.74, ii) S
I and SII leaf like, iii) pecten epipharyngis compris-
ing 3 toothed plates, median with 3 and each lateral
with 8 teeth, iv) premandible with 2 blunt, unequal
apical teeth and 1 small inner tooth and v) pecten
mandibularis unmarked.
Pupa: 1) conical, apically bent cephalic tubercle
bearing subapical seta, ii) pedes spurii B caudolat-
eral on segment II and basolateral on segment I, iii)
tergite I bare, iv) segments II-VIII with 4, 3, 6, 6,
5, 6, 2 pairs of dorsal setae and 3, 2, 3, 3, 3, 4, 4, pairs
of lateral setae, v) caudolateral spur on segment
VII with 9-11 stout spines.
Adult: i) corona with 4 setae, ii) frontal tubercles
absent, 111) haltere light brown bearing 4-5 setae, iv)
scutellum with 18 irregular setae, v) pattern of wing
markings, vi) superior volsella stout, bent bearing
3 basal setae and vii) dark, slender anal point with
an apical rounded knob and seminal capsules open-
ing separately into the vagina.
Remarks. - The adult male and female of this
species were described from West Bengal and Arun-
achal Pradesh, India. It requires mentioning here
that the structural pattern of the teeth of the larval
mentum deviates from that stated by Pinder and
Reiss (1983), but other larval and pupal characters
clearly support its present recombination with the
genus Stictochironomus Kieffer.
Redescription
Adult. - Male: Body 3.93 (3.88-4.04, n = 8) long,
wing 1.62 (1.61-1.64, n = 10) long and 0.52 (0.50-
0.55, n = 10) wide.
Head: Brown. Vertex with 13-15 setae (IV 5-6,
OV 6-7, PO 2). Corona with 4 setae. Clypeus with
24-25 setae, clypeal ratio 0.86. Maxillary palp
brown, ratio of palpomere length I-V: 9 : 15 : 36:
34 : 57; L/W 4.5. Eyes reniform with a dorsal ex-
tension of 0.02 long. Frontal tubercles absent. An-
tenna brown, ratio of flagellomere length I-XI: 15 :
ORTORIORTORIORTORTORAGRIGRAOEMIGS ARS pedicel
ratio 1.15 . CA 0.62; CP 0.99.
165
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Thorax: Dark brown. Antepronotum collar-like
with slight emargination in middle, antepronotals
O. Mesonotum brown. Acrostichals 18 irregularly
biserial, dorsocentrals 19-21 uniserial, prealars 4-5.
Scutellum with 18 irregular setae, postscutellum
dark brown and bare.
Wing (fig. 41): Hyaline. Brachiolum with 1 seta
and 9-11 sensilla campaniformia. R with 17, R,
with 8 and R,,; with 15-16 setae; R,,, meets C at
a distance of 0.108 from R,; RM proximal to FCu.
Wing with a few spots, cells r,;; with 3, cell m1,
with 1 near the fork and cell an with 1 irregular
markings. Squama with 18-19 setae. Haltere light
brown with 4-5 setae. CR 0.95; VR 1-07.
Legs: Yellow to pale brown. Fore tibia with a
blunt scale (fig. 42) bearing 2 long setae. Mid tibia
with single spur 0.021 long, ratio of length of spur
to the apical diameter of mid tibia 7 : 13; hind tibia
with single spur 0.024 long, ratio of length of spur
to the apical diameter of hind tibia 8 : 15. Femur of
fore, mid and hind leg brown; tarsomere V of fore
leg brown, other yellow. Sensilla chactica indistin-
guishable. Pulvilli moderately developed. Propor-
tions and ratio of leg-segments in table 1.
Abdomen: Brown. Hypopygium (fig. 43) with
slender apically knobbed anal point 0.036 long with
3-4 basolateral setae. Gonocoxite with 9-10 setae;
gonostylus stout with rounded apex bearing 12-13
setae along its inner apical margin and 9-10 setae
over it. Superior volsella stout, bent bearing 3 basal
setae; inferior volsella well developed bearing 1
long apical seta and 11-12 incurved setae near apex.
Transverse sternapodeme 0.054, lateral sternapo-
deme 0.108, coxapodeme 0.045 and phallapodeme
(OKO) ons sEIVIO'S75 MERS ND)
Female. - Body 3.49 (3.23-3.58, n = 11) long,
wing 1.62 (1.60-1.65, n = 12) long and 0.48 (0.48-
0.50, n = 12) wide.
Similar to male with usual sex differences. An-
tenna (fig. 44) yellow, flagellomere V brown, ratio
of flagellomere length I-V: 18 : 17 : 20 : 19 : 39,
AR 0.53. Genitalia (fig. 45) with notum 0.093 long.
Coxosternapodeme short. Gonapophysis VIII (fig.
46) divided into long dorsomesal lobe and stout
ventrolateral lobe; apodeme lobe weak. Postgenital
plate V-shaped. Cerci well developed and finely
setose. Seminal capsules equal, almost rounded
0.042 in diameter, ducts of seminal capsules without
any loop, opening separately into the vagina.
Pupa. - Brown. Exuviae pale brown. Body 4.44
(4.34-4.52, n = 8) long in male and 3.89 (3.80-3.96,
n = 8) long in female.
Cephalothorax: Brown. Frontal apotome (fig.
47) with conical, apical bent cephalic tubercles, 0.13
long and 0.10 diameter at base, subapical frontal
seta 0.12 long. Antennal sheath in male (fig. 48)
0.91 long, in female (fig. 49) 0.62 long. Thorax
166
rugose; wing sheath 1.05 long; thoracic horn (fig.
50) with an oval base 0.036 wide and a bunch of
finely branched filaments. 2 pairs of precorneal
setae.
Abdomen (fig. 51): Brown. Pedes spurii A caudo-
lateral on segment V-VIII, not pronounced in seg-
ment VII, pedes spurii B caudolateral on segment
II and basolateral on segment I. Tergite I bare,
tergite II (fig. 52) with broad median patches of
shagreen and a caudal transverse row of 32-41
hooklets; tergites III-IV with a narrow caudal and
median irregular patches of shagreen; tergite V
with median irregular patches of shagreen; tergite
VI with a basal, narrow median and caudal trans-
verse patches of shagreen, tergites VII-IX with 2
basal patches of shagreen. Segments II-VIII with 4,
3, 6, 6, 5, 6, 2 pairs of dorsal setae and segments II-
VIII with 3, 2, 3, 3, 3, 4, 4 pairs of lateral setae, on
segment V-VIII filamentous; segment VIII with 1
pair of caudolateral spurs (fig. 55) 0.18 long, each
with 9-11 stout spines. Anal fin (fig. 51) 0.23 long
with numerous filamentous setae. Genital sac in
male (fig. 53) 0.20 long, in female (fig. 54) 0.22
long. G/F 0.87 in male and 0.96 in female.
Fourth instar larva. - Colouration dark red. Ex-
uviae pale. Head capsule, claws of anterior and
posterior parapods, and anal setae brown. Body 6.41
(6.29-6.55, n = 12) long.
Head: Brown. Occipital margin dark brown.
Ventral head capsule (fig. 56) 0.27 long and 0.20
wide. Two eye spots, dorsal one slightly larger than
ventral. Antenna (fig. 57) six-segmented, basal an-
tennal segment 0.03 long and 0.012 wide, with a
ring organ 0.003 in diameter, distance to ring organ
from base 0.018 long; blade of basal antennal seg-
ment 0.054 long; accessory blade 0.012 long; Lau-
terborn organ two, first on antennal segment II
being 0.012 long and second on segment III being
0.003 long; ratio of antennal segment length I-V:
10:3:4:1.5 :4:1; AR 0.74. Labral lamella rod
shaped with median pouch; S I (fig. 58) leaf like
with serrated margin 0.021 long, between the bases
of two S I 0.016 long; S II also leaf like 0.015 long;
S III short with prominent base; S IV simple; S V
minute; 4 chaetae; 2 spinulae; 4 chaetulae laterales;
chaetulae basales absent. Pecten epipharyngis (fig.
59) comprising 3 toothed plates, median with 3 and
each lateral with 8 teeth. Premandible (fig. 60)
0.054 long with 2 unequal, blunt apical teeth and 1
small inner tooth, premandibular brush dense.
Mandible (fig. 61) 0.09 long with 1 apical and 3
inner teeth, first one smaller than others; seta sub-
dentales 0.027 long; seta interna with 2 main plu-
mose branches, one longer than the other; 2 seta
externa; pecten mandibularis unmarked; ring organ
0.004 in diameter, very close to the base. Maxilla
with blade like anterior chaeta; roughly pointed
lacinial chaetae; antaxial seta and paraxial seta well
developed; sensillum basiconica 1; pecten galearis
distinct; maxillary palp 3 segmented; first segment
of maxillary palp 0.014 long, 0.012 wide, ring organ
0.082 in diameter, 4 setae maxillaris, SM, and SM,
close to the palpiger, SM; and SM, close together
on the sclerite of cardo. Prementohypopharyngeal
complex (fig. 62) with prementum 0.021 wide, me-
dian lamella distinct, paramedian lamella indis-
dinct, 4 sensilla and 6-7 chaetulae. Mentum (fig. 63)
0.048 long and 0.075 wide with a trifid median
tooth 0.018 long and 0.021 wide; 6 pairs of lateral
teeth, size gradually decreases from Ist lateral to the
4th, 5th long 6th short. Ventromental plate (fig. 64)
fan-shaped 0.057 wide with distinct rays ending
before the margin. V/M 2.71.
Abdomen: Dark red. Procercus 0.018 long, 0.027
wide at base, each with 8 anal setae 0.31 long and
2 short lateral setae. 2 supraanal setae, 0.29 long;
Sa/An 0.93. Anterior parapods with numerous
weakly sclerotised claws. Posterior parapods (fig.
65) 0.09 long with 15-16 well sclerotised variable
claws (fig. 66). Anal tubules (fig. 65) conical 0.05
long.
40. Xenochironomus flaviventris (Kieffer)
Chironomus flaviventris Kieffer, 1911: 139.
Xenochironomus flaviventris (Kieffer): Chaudhuri &
Guha 1987: 29.
Material examined. - 1 8, Farakka, 29 September 1986;
1 & Hooghly, 4 September 1987; 4 3, 6 9, Naihati, 16
September 1987; 1 8 paratype (Z.S.I. Regn. No.
9731/19), India, Orissa, Puri, 20-21 January 1908, leg. N.
Anandale.
Distribution. — India.
Differential diagnosis. — i) frontal tubercles ab-
sent, 11) scutellum with 12-13 setae in a row, iii)
brachiolum with 3 setae, iv) colour pattern of leg,
v) tergites with dark stripes, vi) anal point short,
broad and dark brown, vii) superior volsella lobe
like, setose and viii) inferior volsella long and dark
brown.
Remarks. — The adult male was described by
Kieffer (1911) in the genus Chironomus Meigen
from Puri, India. On examining a paratype speci-
men from NZC, Calcutta, Chaudhuri and Guha
(1987) placed this species in the genus Xenochtro-
nomus Kieffer. Reexamination of this paratype and
a comparison with specimens from rice fields of
Gangatic West Bengal confirm its generic position.
41. Cladotanytarsus conversus (Johannsen)
Tanytarsus conversus Johannsen, 1932: 543.
Cladotanytarsus conversus; Sublette & Sublette 1973: 416.
Material examined. - 8 8, Kalyani, 25 October 1986.
Distribution. - South Sumatra and India (present
record).
CHAUDHURI & CHATTOPADHYAY: Chironomids
Differential diagnosis. — i) frontal tubercles
small, 11) scutellum with 8-10 irregular setae, iii)
pulvilli very small, iv) tergites I-VII brown at lateral
and caudal margin and v) anal point short without
punctures.
42. Cladotanytarsus gloveri Ghosh & Chaudhuri
Cladotanytarsus glovert Ghosh & Chaudhuri, 1983: 126.
Material examined. - 4 8, Burdwan, 14 September
1986; 3 8, Chinsura, 6 February 1987.
Distribution. — India.
Differential diagnosis. — i) frontal tubercles small,
it) scutellum with 6 setae, iii) haltere with 4-7 setae,
iv) tergites I-VI brown at their caudal margin and
v) anal point with 6-8 pucntures.
43. Cladotanytarsus multispinulus Guha, Das,
Chaudhuri & Choudhuri
Cladotanytarsus multispinulus Guha, Das, Chaudhuri &
Choudhuri, 1985: 31.
Material examined. - 9 8, Kalyani, 17 February 1987;
2 à, Uttarpara, 24 February 1987; 3 6 Burdwan, 6 Sep-
tember 1987.
Distribution. — India.
Differential diagnosis. — i) frontal tubercles ab-
sent, ii) scutellum with 4 setae, iii) tergites I-V
whitish with a yellowish apical band, tergites VI-
VIII dark yellow and iv) anal point small with
numerous spinules.
44. Tanytarsus bifurcus Freeman
Tanytarsus bifurcus Freeman, 1958: 337; Freeman &
Cranston 1980: 221; Chaudhuri et al. 1984: 33; Chaud-
huri & Guha 1987: 29.
Material examined. - 5 8, Farakka, 29 September, 1986.
Distribution. - Africa (Upper Volta) and India.
Differential diagnosis. — i) corona with setae, ii)
scutellum with 6 setae, iii) haltere bare and iv) anal
point stout with bifurcated apex.
45. Tanytarsus commoni Glover
Tanytarsus commoni Glover, 1973: 458.
Tanytarsus tamakutibasi Sasa, 1983: 22. Syn. n.
Material examined. - 6 8, Kalyani, 25 October 1986; 1
8 paratype (Aust. Nat. Ins. Coll., Australia), Clyde Mtn.,
Eastern foothills, 40 ft. N.S.W., 15 August 1967, leg. Z.
Liepa.
Distribution. - Australia, Japan and India (pres-
ent record).
Differential diagnosis. — i) frontal tubercles oval,
ii) prescutellar 1, iii) pulvilli absent, iv) anal point
167
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
with 4 lateral setae and a subterminal oval lobe
bearing 5 spinulae and v) median volsella small
brush like.
Remarks. - A study of a male paratype from
ANIC, Australia and a comparison with the present
ones from rice fields confirm the conspecificity.
46. Tanytarus fuscimarginalis Chaudhuri, Guha
& Ghosh
Tanytarsus fuscimarginalis Chaudhuri, Guha & Ghosh,
1984: 33-34; Chaudhuri & Guha 1987: 30.
Material examined. - 4 8, Kalyani, 10 February 1986;
5 6, Uttarpara, 24 February 1987.
Distribution. — India.
Differential diagnosis. — i) scutellum with 4 setae,
ii) wing white with yellowish tinge, ii) haltere se-
tose, iv) abdominal tergites brown at the caudal and
lateral margins and v) hypopygium with anal point
bearing 5-7 punctures in a row.
47. Tanytarsus vinculus Chaudhuri, Guha &
Ghosh
Tanytarsus vinculus Chaudhuri, Guha & Ghosh, 1984: 34;
Chaudhuri & Guha 1987: 30.
Material examined. - 7 g, Howrah, 4 September 1987.
Distribution. — India.
Differential diagnosis. — i) corona bare, ii) frontal
tubercles present, iii) haltere bare, iv) tergite I with
a basal brown band, others with similar band at the
caudal margin and v) hypopygium with anal point
bearing 4 punctures in a row; median volsella
brush-like with 8-10 unbranched setae.
48. Clinotanypus fuscosignatus (Kieffer)
Procladius fuscosignatus Kieffer, 1910: 218.
Clinotanypus fuscosignatus; Sublette & Sublette 1973:
390.
Material examined. — 5 larvae, Howrah, 4 September
1987; 5 larvae, 8 @ and 8 9 pupae, 6 @ and 5 © exuviae,
7 6, 10 9, all reared; 2 g, Bally, 8 March 1987; 1 4,
Burdwan, 6 September 1987.
Distribution. - Bangladesh and India.
Differential diagnosis. - Larva: i) high value of
AR (15.2), ii) chaetulae laterales and chaetulae ba-
sales indistinct, ii) middle two teeth of ligula small
and equal, iv) paraligula with 3 points on inner side
and 2 points on outer side and v) mentum with 14-
15 dorsomental teeth on each side in two rows.
Pupa: i) frontal apotome without cephalic tuber-
cles and frontal setae, ii) thoracic horn with short
neck and broad plastron plate, iii) segments I-VI]
with 2, 1, 3, 3, 3, 3, 6 pairs of dorsal setae, iv) lateral
setae on segments VII and VIII simple and v) G/F
0.43 in male and 0.39 in female.
168
Adult: i) corona with 4 setae, ii) frontal tubercles
absent, iii) mesonotum with 6 bands, iv) scutellum
with 40-42 long setae in two rows and numerous
small setae, v) pattern of wing markings, vi) band-
ing pattern of abdomen and vii) tibial spurs with 5
teeth.
Remarks. - The adults of both sexes of Procladius
fuscosignatus were first described by Kieffer (1910)
from Calcutta, India. Sublette & Sublette (1973)
transferred it to Clinotanypus Kieffer. Study of type
specimens from NZC, Calcutta and comparison
with material reared in the laboratory confirm the
validity of the species and the combination sug-
gested by Sublette & Sublette (1973).
Redescription
Adult. - Male: Body 4.07 (3.99-4.14, n= 10) long,
wing 2.25 (2.23-2.27, n = 10) long and 0.91 (0.90-
0.92, n= 10) wide.
Head: Reddish brown. Vertex with 27-29 setae
(IV 3-4, OV 6, PO 18-19). Corona with 4 setae.
Clypeus with 14-16 setae, clypeal ratio 1.66. Max-
illary palp pale brown, ratio of palpomere length I-
V:6 :10 :40 :5 : 70; L/W 3.07. Eyes reniform with
a dorsal extension of 0.08 long. Frontal tubercles
absent. Antenna brown, ratio of flagellomere
length E.G 14 39) S10) 0) 89) 3 te}-2 10) s Be NSR:
8 : 8 : 260 : 50; AR 2.81; pedicel ratio 1.0. CA 0.47;
GP REE
Thorax: Shiny light brown. Antepronotum well
developed, antepronotals 10. Mesonotum with 3
pairs of bands in antero-median, postero-lateral
and prescutellar region. Acrostichals 20-21 irregu-
larly biserial, dorsocentrals 21-23 irregularly bise-
rial, humerals 11, prescutellars 30-32, prealars 23-
30 in a staggered row. Scutellum shiny with 40-42
setae in two rows at the posterior margin in addi-
tion to numerous irregular small setae, postscutel-
lum dark brown except the upper end bearing 30-
ID SEE,
Wing (fig. 67): Smoky and distal third hyaline.
Brachiolum with 3-4 setae and 26-30 sensilla com-
paniformia. Costa extended a little being 0.08 long,
R, shortly bifurcated at the tip; RM thickened,
moderately oblique; MCu proximal to RM; RM
slightly proximal to FCu. Wing with characteristic
bands and spots; dark brown transverse band across
distal half from R, to twothird of cell m beng
interrupted in cell r,;;, a large square brown spot
over RM. Squama with 29-31 setae, haltere pale
with 6-8 setae. CR 0.97; VR 1.07.
Legs: Yellow. Femora of all legs and fore tibia
with a basal and apical, mid tibia with only basal and
hind tibia with basal and apical dark brown bands.
Fore tibia with single spur (fig. 68) 0.054 long
having 9-10 small weak lateral teeth, ratio of length
of spur to the apical diameter of fore tibia 18 : 25;
spurs of mid tibia (fig. 69) unequal, 0.042 and 0.051
long, each having 5 lateral teeth, ratio of length of
spurs to the apical diameter of mid tibia 14 : 24 and
17 : 24; spurs of hind tibia also unequal, 0.048 and
0.081 long, each with 5 lateral teeth, ratio of length
of spurs to the apical diameter of hind tibia 16 : 30
and 27 : 30. Hind tibial comb with 13 setae each.
Tarsomeres IV-V entirely brown. Proportions and
ratios of leg-segments in table 1.
Abdomen (fig. 70): Yellowish brown. Tergites II-
VII with transverse brown bands and tergites V and
VIII with very narrow transverse brown bands.
Tergite IX with 22-23 setae. Hypopygium (fig. 71)
with triangular anal point. Gonocoxite 0.24 long,
broad proximally and narrow distally; gonostylus
(fig. 72) 0.15 long, distal part with 5 setae, curved
at right angle to the base and with a subapical tooth.
EIRWIEG gE; 2:66:
Female. - Body 2.92 (2.65-3.24, n= 4) long, wing
2.19 (2.03-2.37, n = 6) long and 0.86 (0.83-0.98, n
= 6) wide.
Similar to male with usual sex differences. An-
tenna brown with 14 flagellomeres, flagellomere II
almost twice as long as broad, flagellomere III-XIV
globular, flagellomere XV elongated, somewhat el-
liptical. Thoracic and leg colouration darker than in
male. Genitalia (fig. 73) with notum 0.22 long.
Coxosternapodeme curved, thickened and broad
proximally. Gonapophysis VII triangular. Post-
genital plate semicircular. Cerci 0.105 long and
finely setose. Seminal capsules 3, approximately
equal, baloon shaped, ducts of seminal capsules
without any loop, opening separately into the vag-
ina.
Pupa. - Brown. Exuviae pale. Body 4.72 (4.66-
4.79, n = 8) long in male and 4.97 (4.91-5.07, n =
8) long in female.
Cephalothorax: Dark brown. Frontal apotome
(fig. 74) without cephalic tubercles and frontal se-
tae. Antennal sheath in male (fig. 75) 1.09 long, in
female (fig. 76) 0.63 long. Thorax rugose; wing
sheath 1.69 long; thoracic horn (fig. 77) 0.4 long
with short neck and broad plastron plate, having
straight basal edge.
Abdomen (fig. 78): Brown. Segments II-VIII
with 1, 1, 1, 1, 1, 7, 5 pairs of lateral setae and
segments I-VII with 2, 1, 3, 3, 3, 3, 6 pairs of dorsal
setae; all setae branched except lateral long setae on
segments VII and VIII. Anal fin (fig. 78) 0.71 long
with 2 pairs of long filamentous setae. Genital sac
in male (fig. 79) 0.31 long, in female (fig. 80) 0.28
long; G/F 0.43 in male and 0.39 in female.
Fourth instar larva. - Colouration light brown,
exuviae palen. Head capsule, claws of anterior and
posterior parapods, and anal setae brown. Body 7.55
(7.32-7.67, n = 10) long.
Head: Brown. Occipital margin pale. Ventral
head capsule (fig. 81) 0.67 long. One eye spor, re-
CHAUDHURI & CHATTOPADHYAY: Chironomids
niform. Antenna (fig. 82, 83) four-segmented, basal
antennal segment 0.43 long with a ring organ 0.004
in diameter, distance to ring organ from base 0.39;
blade of basal antennal segment 0.036 long; acces-
sory blade 0.015 long; blade of antennal segment II
0.021 long; Lauterborn organs 2, very small; ratio
of antennal segment length I-IV: 144 : 7: 1.5 : 1;
AR 15.2. S I simple 0.018 long, between the bases
of two S 10.021 long; STI-HI simple; S TV indistinct;
2 chaetae; 3-4 spinulae; chaetulae laterales and
chaetulae basales indistinct. Mandible (fig. 83)
0.129 long with expanded base on inner side, apical
tooth dark and strongly hooked, lateral teeth 4; first
lateral tooth large, pointed apically; others small;
seta subdentalis distinct; 3 seta externa; ring organ
0.003 in diameter, distance of ring organ from base
0.006. Maxilla (figs. 85, 86) developed, maxillary
palp 3 segmented, first segment of maxillary palp
0.066 long with a ring organ 0.003 in diameter;
segment II hyaline and 4-5 sensilla at its apex.
Prementohypopharyngeal complex well deve-
loped; ligula (fig. 87) with 6 teeth, middle two small
and equal; paraligula (fig. 87) with 3 points on
inner side and 2 points on outer side. Mentum (fig.
88) with 14-15 light brown dorsomental teeth on
each side laterally in double row; M appendage (fig.
88) with distinct, granulose pseudoradula, labial
vesicles absent.
Abdomen: Light brown. Procercus 0.165 long
and 0.054 wide, each with 15-16 anal setae 0.658
long and 2 small lateral setae. 2 supraanal setae 0.53
long; Sa/An 0.808. Anterior parapods with numer-
ous weakly sclerotised claws. Posterior parapods
(fig. 89) 0.434 long with 14-15 sclerotised variable
claws (fig. 90). Anal tubules (fig. 89) conical, 0.21
long.
49. Procladius noctivagus (Kieffer)
Procladius noctivagus; Freeman 1955c: 59; Freeman &
Cranston 1980: 177; Chaudhuri & Debnath 1983: 121
(as subgenus).
Trichotanypus niloticus Kieffer, 1923: 189.
Trichotanypus nilicola Kieffer, 1925: 309.
Material examined. — 5 larvae, Kalyani 10 February
1986; 3 larvae, 9 & and 9 © pupae, 4 & and 6 Q exuviae,
7 6,8 9, all reared; 1 g, Burdwan, 29 January 1986.
Distribution. - Chad, Egypt, India, Niger, Nige-
ria and Sudan.
Differential diagnosis. — Larva: i) AR 4.0, ii)
mandible with 2 minute elevations at the inner
apical margin, iii) ligula with 5 dark teeth, middle
one shortest, iv) paraligula with 4 points on inner
side and 7 points on outer side, v) mentum with 8
dorsomental teeth on each side and vi) M appen-
dage with bulb like labial vesicles.
Pupa: 1) thoracic horn linear with oval plastron
plate, ii) tergites I and III bare, iii) segments I-VIII
169
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
with 3, 2, 2, 6, 5, 5, 5, 2 pairs of dorsal setae and iv) G/F
0.44 in male and 0.4 in female.
Adult: i) frontal tubercles absent, ii) scutellum with 20-
24 irregular setae, iii) haltere yellow with 6-7 setae, iv)
tibial spurs with 4 teeth, v) hind tibialcomb with 11 setae,
vi) banding patterns of abdomen and vii) gonostylus
small, heel prominent and having a sharp tooth.
Remarks. - The analysis of characters of both imma-
tures and adults reconfirm its systematic position.
Redescription
Adult. - Male: Body 3.61 (3.51-3.68, n = 8) long,
wing 1.57 (1.55-1.58, n = 8) long and 0.53 (0.53-
0.55, n = 8) wide.
Head: Brown. Vertex with 25-26 setae (IV 3-4,
OV 12, PO 10). Corona with 4 setae. Clypeus with
16 setae, clypeal ratio 0.57. Maxillary palp dark
brown, ratio of palpomere length I-V:4 :5 :7 :12:
20; L/W 2.33. Eyes reniform with dorsal extension
of 0.09. Frontal tubercles absent. Antenna brown,
ratio of flagellomere length I-XIV:2:3:4:5:5:
32603023028 28 3 11028 285 AUR 22138 pediceltratio
RORCMOPAICPIDO!
Thorax: Yellowish brown. Antepronotum with a
dorsal emargination in middle, antepronotals 6.
Mesonotum with 3 light brown vittae. Acrostichals
36-40 irregularly biserial and diverging posteriorly,
dorsocentrals 15 uniserial, humerals 5, prescutellars
6, prealars 10. Scutellum with 20-24 irregular setae,
postscutellum dark brown and bare.
Wing ((fig. 91): Wing membrane clothed with
macrotrichia. Brachiolum with 2 setae and 19-21
sensilla campaniformia. Costa extended, extension
0.12 long. Wing with spots at the fork of R, cell rs,
cell m, cell m;,, and cell an; RM clouded and base
of MCu thickened, RM oblique, MCu below RM.
Squama with 23-25 setae. Haltere yellow with 6-7
setae. CR 0.95; VR 1.45.
Legs: Yellowish brown. Femora yellow, apex of
tibia dark. Fore tibia with a spur (fig. 92) 0.062 long
with 4 lateral teeth, ratio of length of spur to the
apical diameter of fore tibia 12 : 14; spurs of mid
tibia (fig. 93) subequal, 0.031 and 0.04 long, each
with 4 lateral teeth, ratio of length of spurs to the
apical diameter of mid tibia 8 : 13 and 10 : 13; spurs
of hind tibia unequal, 0.03 and 0.05 long, each with
4 lateral teeth, ratio of length of spurs to the apical
diameter of hind tibia 8 : 13 and 10 : 13; spurs of
hind tibia unequal, 0.03 and 0.05 long, each with 4
lateral teeth, ratio of length of spurs to the apical
diameter of hind tibia 8 : 16 and 15 : 16. Apex of
tarsomeres I-III narrowly darkened, tarsomeres IV-
V uniformly brown. Mid and hind tarsomeres I-II
with single spur 0.031 long. Hind tibia comb (fig.
94) with 11 setae. Pulvilli absent. Proportions and
ratios of leg-segments in table 1.
Abdomen (fig. 95): Yellow. Tergites with a basal
transverse dark brown band, tergites V-VIII mostly
dark with pale caudal area. Hypopygium (fig. 96)
with scale like anal point. Gonocoxite stumpy,
170
broad at the base and narrow distally; gonostylus
small, heel prominent and having a sharp tooth.
Apodemes well developed. HR 2.44; HV 3.58.
Female. - Body 2.56 (2.51-3.02, n= 8) long, wing
1.71 (1.68-1.74, n = 8) long and 0.66 (0.62-0.68, n
= 8) wide.
Similar to male with usual sex differences. An-
tenna brown, ratio of flagellomere length I-XIII :
© 26 26 2626 26 26 2625 392527 226 ANR 0,37
Genitalia (fig. 97) with notum 0.18 long. Coxo-
sternapodeme curved. Gonapophysis VIII triangu-
lar with a fold. Postgenital plate developed. Cerci
finely setose. Seminal capsules 3, approximately
equal, ovoid 0.07 long by 0.06 wide; duct of seminal
capsules without any loop, opening separately into
the vagina.
Pupa. - Brown. Exuviae white. Body 3.19 (3.09-
3.26, n = 9) long in male and 3.49 (3.39-3.51, n=
9) in female.
Cephalothorax: Brown. Frontal apotome (fig.
98) without cephalic tubercles and frontal seta. An-
tennal sheath in male (fig. 99) 0.74 long, in female
(fig. 100) 0.48 long. Thorax rugose; wing sheath
0.89 long; thoracic horn (fig. 101) linear, 0.28 long
with oval plastron plate and distinct neck.
Abdomen (fig. 102): Brownish yellow. Tergites I
and HI bare; tergite II and tergites IV-VIII with
median shagreen. Segments I-VII with 3, 2, 2, 6, 5,
5,5, 2 pairs of dorsal setae and segments II-VII with
2, 2, 1, 1, 1, 4, 4 lateral setae, on segmentous VII-
VIII filamentous. Anal fin (fig. 102) 0.35 long with
2 pairs of filamentous setae. Genital sac in male (fig.
103) 0.15 long, in female (fig. 104) 0.14 long, G/F
0.44 in male and 0.4 in female.
Fourth instar larva. - Brownish yellow. Exuviae
white. Head capsule, claws of anterior and posterior
parapods and anal setae brown. Body 4.61 (4.54-
4.69, n = 8) long.
Head: Brown. Occipital margin dark brown.
Ventral head capsule (fig. 105) 0.39 long and 0.38
wide. One eye spot, almost reniform. Antenna (fig.
106) four-segmented, basal antennal segment 0.096
long and 0.21 wide at base with a ring organ 0.004
in diameter, distance to ring organ from base 0.069;
blade of basal antennal segment 0.018 long; acces-
sory blade 0.004 long; blade of antennal segment II
0.0015 long, 1 pair of minute Lauterborn organ;
ratio of antennal segment length I-IV: 32: 5.5:
1.5 : 1; AR 4.0. Labral lamella more or less trian-
gular. S I blade like 0.015 long; S II tubular with an
apical setae, S III and S IV minute; 1-2 chaetae; 2
spinulae; chaetulae laterales and chaetulae basales
absent. Mandible (fig. 107) 0.111 long with 1 dark
apical tooth, 1 prominent basal tooth and 1 conical
tooth at the base of apical tooth; seta subdentalis
0.015 long; 3 seta externa; inner margin with 2
minute elevations at the apical portion. Maxilla
(fig. 108) developed, maxillary palp 3 0.042 long
with a ring organ 0.0015 in diameter and 5-6 sen-
silla at its apex; 2 setae maxillaris; SM, and SM,
close to the stipes.
Prementohypopharyngeal complex well deve-
loped; ligula (fig. 109) with 5 dark teeth, middle one
shortest; paraligula (fig. 109) with 4 points on
inner side and 7 points on outer side. Mentum (fig.
110) with 8 brown dorsomental teeth on each side,
outer one smallest; M appendage (fig. 110) with
bulb like labial vesicles, pseudoradula distinct.
Abdomen: Brownish yellow. Procercus 0.14 long
and 0.039 wide, each with 14-15 anal setae 0.532
long and 2 small lateral setae. 2 supraanal setae
0.392 long, Sa/An 0.74. Anterior parapods with
numerous weekly sclerotised claws, posterior pa-
rapods (fig. 111) 0.36 long with 14-15 sclerotised
variable claws (fig. 112). Anal tubules (fig. 111)
conical 0.25 long.
50. Tanypus bilobatus (Kieffer)
Procladius bilobatus Kieffer, 1913b: 155.
Tanypus bilobatus; Chaudhuri, Nandi & Ghosh 1983: 122-
133; Chaudhuri & Guha 1987: 25.
Material examined. — 6 larvae, Burdwan, 30 May 1986;
9 larvae, 6 & and 6 9 pupae, 6 8 and 5 9 exuviae, 5 Á, 5
Q, all reared; 3 8, Naihati, 16 September 1987; 4 6,
Hooghly, 4 September 1987.
Distribution. — India.
Differential diagnosis. — 1) larval antenna only
about one-third of the head length, middle tooth of
ligula as long as those of lateral, number of teeth in
paraligula and shape of mentum, ii) pupa with
biconvex thoracic horn and transparent frontal ap-
otome without cephalic tubercle, frontal setae 1
pair, iii) adult with setose haltere, rounded anal
lobe, MCu proximal to FCu distance between MCu
and base of FCu less than onethird as long as Cu,,
tarsomere V uniformly brown and wing with nu-
merous spots or bands.
Remars. - On examination of the type specimens
present at the NZC, Calcutta and correlating them
with the collected and reared material, Chaudhuri et
al. (1983) recombined this species with the genus
Tanypus.
51. Tanypus grandis Chaudhuri, Das & Debnath
Tanypus grandis Chaudhuri, Das & Debnath, 1984: 100;
Chaudhuri & Guha 1987: 25.
Material examined. — 10 8, Kalyani, 21 August 1986.
Distribution. — India.
Differential diagnosis. — 1) large size, ii) linear
brown streak near the base of cell r and m, iii) spots
near the margin of cell m beyond MCu, cell m; ;,
CHAUDHURI & CHATTOPADHYAY: Chironomids
with two rounded spots, near its distal margin, iv)
tergites with dorsomedian dark patches, v) chaeto-
taxy of thorax and vi) gonostylus with prominent
carina.
52. Tanypus lucidus Chaudhuri, Das & Debnath
Tanypus lucidus Chaudhuri, Das & Debnath, 1984: 103
Chaudhuri & Guha 1987: 25.
Material examined. - 4 @, Bally, 13 February 1986; 8 3,
Burdwan 7 September 1987.
Distribution. - India.
Differential diagnosis. — i) scutellum with 30-32
setae, ii) brachiolum with 1 seta, iii) squama with
31 setae, iv) tibia uniformly brown and hypopygium
without anal point or with membranous structure,
v) hind tibial comb with 5 setae and vi) cell r
without any markings.
53. Tanypus tenebrosus Chaudhuri, Das &
Debnath
Tanypus tenebrosus Chaudhuri, Das & Debnath, 1984:
105; Chaudhuri & Guha 1987: 25.
Material examined. - 5 8, Naihati, 16 September 1987.
Distribution. — India.
Differential diagnosis. — i) brachiolum with 2
setae, ii) haltere brown with 4 setae, iii) tarsomeres
uniformly brown, iv) hind tibial comb with 9 setae,
v) tergites I-V with mid lateral stripe, tergites VI-
VIII with only middle stripe and vi) gonostylus
small, bent ending in an apical tooth.
BIOLOGY
Almost all the rice fields of West Bengal are
polluted. Chironomid larvae accumulate either by
eggs deposited there or are carried to the field with
incoming water from various sources. At the begin-
ning of the rice season, the fields are muddy and
studded with larval tubes of chironomids belonging
to the genus Chironomus. In the middle of the
season larvae of Chironomus were seen to live there
in association with algal mats. All the species of
chironomid larvae, except for the predaceous forms
belonging to the genera Clinotanypus, Tanypus and
Procladius, constructed tubular houses. The tubes
were mostly ‘U’-shaped or ‘S’-shaped in Chirono-
mus and Kiefferulus and sometimes tunnel-like in
the mud. The houses of Polypedilum and Stictoch:-
ronomus were usually slender and ‘U’-shaped, but
conical tubes appeared at lower depths than in Chz-
ronomus. The nests of Dicrotendipes were smaller
than those of Chironomus. The tubes built by the
larvae of Tanytarsus were distinguished into two
types, long cylindrical, or upright tubes over or just
beneath the mud surface in winter and branched in
171
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 2. Duration of life history stages in summer (32 + 2.5
°C) and winter (24 + 2.5 °C); a = summer, b = winter.
Species Egg
(hrs)
Chironomus samoensis a 34-50
b 52-74
Dicrotendipes pelochloris a 30-54
b 36-71
Stictochironomus obscurum a 30-42
b 46-73
Clinotanypus fuscosignatus a -
b 59-90
Procladius noctivagus a 52-84
b 64-88
the summer and remaining in the deeper layer of
substrates.
It was observed that the larvae of Chironomus
samoensis and Kiefferulus barbatitarsis prefer to
remain in clumped conditions in association with
the roots of the rice plants. It seems interesting that
the larvae of Cryptochironomus judicius made their
abodes on the side walls of the rice fields, whereas
there was no trace of the larvae within the rice
fields. It may be thought that Tanypus larvae were
dominant in the early crop season whereas the
larvae of Clinotanypus dominated in the late sea-
son. In contrast to the observations of Ping (1917)
and Darby (1962), larvae of K. barbatitarsis sur-
vived in most cases for almost one month out of
water, but the remaining larvae died after four to
five days under such conditions.
The present study details the life history of 5
species: Chironomus samoensis Edwards, Dicro-
tendipes pelochloris (Kieffer), Stictochironomus
obscurus (Guha & Chaudhuri), Clinotanypus fusco-
signatus (Kieffer) and Procladius noctivagus
(Kieffer). The duration of egg, larva, pupa and
imago in both summer (32 + 2.5 °C temperature
and 51-76% humidity) and winter (24.5 + 2.5 °C
temperature and 20-48% humidity) are shown in
table 2.
EMERGENCE AND SEX RATIO
Emergence of adult chirnomids was studied in
the laboratory in both summer (32 + 2.5 °C) and
winter (24.5 + 2.5 °C). The emergence period ap-
peared to be related to the duration of the larval
period. Chironomus samoensis emerged in the ev-
ening hours, i.e. at dusk. The emergence period in
winter was 1-2 hours earlier than in summer, which
might be due to the change in the time of sunset.
Dicrotendipes pelochloris and Stictochironomus
obscurus emerged mostly throughout the day ie.
172
Larval Pupa Adult Total
duration (hrs) (days) (days)
(days)
16-25 24-28 2-35 20.42-31.75
24-36 27-38 3-4.5 30.29-45.17
20-29 24-48 2-4 24.25-37.25
25-40 34-56 3-4 30.92-49.29
20-26 21-29 2-3 24.12-31.96
21-28 26-36 3-4 27.00-36.54
64-72 51-65 3-4 71.58-82.46
45-50 46-56 1-3 50.08-59.25
55-68 59-75 2-3 62.12-77.79
dawn and dusk in summer. In winter maximum
emergence occurred during the mid day except for
D. pelochloris where it also continued in the after-
noon, up to the early hours of darkness during
several days of observations. Procladius noctivagus
exhibited a bimodal pattern in summer, emerging
in dawn and dusk with a few hours (1-2) earlier in
winter. In winter, the emergence of Clinotanypus
fuscosignatus began 2-3 hours prior to dawn and
dusk with a peak of 1-1.5 hours after sunset.
It is evident here that the adult emergence of
most chironomids showed an approximately diel
periodicity and maximum emergence occurred
mainly between late afternoon and early hours of
darkness. The emergence pulse tended to be higher
near the middle of the total emergence period.
Sex ratio in laboratory emergences (table 3)
The sex ratio of adults reared from egg masses in
the laboratory was expressed in percentages of
males, or females. The percentage of males was
higher in Chironomus samoensis and Stictochiro-
nomus obscurus. On the other hand, females pre-
dominated in Dicrotendipes pelochloris, Clinotany-
pus fuscosignatus and Procladius noctivagus. Of the
five species of chironomids studied here, the high-
est percentage of males was observed in Stictochi-
ronomus obscurus (65.55%) and of females in C4-
notanypus fuscosignatus (67.07%).
DISCUSSION
Investigations represented in this paper com-
prise morphological diagnoses of 53 species of chir-
onomids found in the rice paddies of West Bengal,
India. Of these, 40 species are recorded as adults
only and 13 including immature stages. Prior to this
work, most of these were recorded from various
places in India other than rice paddies. Further 10
CHAUDHURI & CHATTOPADHYAY: Chironomids
Table 3. Numbers of larvae, pupae and adults reared from egg masses and sex ratios of emerged adults.
Species No. of No. Larvae Percen- Pupae Percen- Adults emerged Percen- Sex ratios
egg of hat- tage of obt- tage of Mr Eee of of adults
masses eggs ched eggs ained larvae pupae (d: 9)
hatched pupated hatched
Chironomus
Samoensis 5 1024 799 78.03 293 36.67 109 61 58.02 64.12:35.88
Dicrotendipes
pelochloris 4 682 374 54.84 168 44.92 49 68 69.64 41.88:58.12
Stictochironomus
obscurum 5 1665 1199 72.01 423 35.28 177 95) 63.83 65.56:34.44
Clinotanypus
fuscosignatus 5 651 377 57.91 103 27.32 Di, DO 79.61 32.93:67.07
Procladius
noctivagus 5 901 602 66.81 225 37.37 54 76 57.78 41.54:58.46
species were previously reported from Thailand
(Hashimoto et al. 1981), six from Japan (Sasa &
Kikuchi 1986) and one from North America
(Darby 1962).
The life cycles of chironomids described here,
present interesting features of the biology of the
midges. The present work is the first attempt to
study the life cycle of the following five oriental
species from rice paddies: Chironomus samoensis
Edwards, Dicrotendipes pelochloris Kieffer, Sticto-
chironomus obscurus (Guha & Chaudhuri), Clino-
tanypus fuscosignatus (Kieffer) and Procladius noc-
tivagus Kieffer. The duration of each stage was
generally neglected in earlier works. It can only be
conveniently determined in the laboratory, as did
Hilsenhoff (1966). Grodhaus (1963) reported the
life cycle of Chironomus tentans Fabricius on the
basis of information from Sadler (1935). The dura-
tion of each stage is temperature-dependent and
related to the overall length of the life cycle (Oliver
1971). The time required for hatching of eggs was
recorded as 2.5 days at 24 °C in Chironomus atrella
(Townes) by Anderson & Hitchcock (1968), 4 days
at 20 °C in C. zealandicus Hudson by Forsyth
(1971) and 4-5 days at 18 °C in some other species
(Mundie 1956). Similarly the duration was recorded
to be 3-4 days at 18 °C in Einfeldia synchrona
Oliver (Danks 1971) and 2-2.5 days and 3-3.5 days
at 32 + 2.5 °C and 24.5 + 3 °C respectively in
Tanypus bilobatus (Kieffer) by Chaudhuri et al.
(1983). Recently, Chaudhuri & Ghosh (1986) re-
ported that the duration of egg stage required 40-
60 hours and 48-72 hours in Kiefferulus barbatitar-
sas and 24-36 hours and 48-69 hours in K. calligaster
atemperatureskofns reen? DR Gand is 37€
respectively. Chattopadhyay et al. (1988) stated the
duration in the case of Polypedilum nubifer to be
28-40 hours in summer (32 + 2.5 °C) and 42-72
hours in winter (24 + 2.5 °C). It may therefore be
inferred that the hatching period varies with
temperature, being shorter with the rise of temper-
ature in summer and longer with the fall of temper-
ature in winter.
The duration of the larval stages of four species,
both in summer and winter and one species in
winter only are presented in table 2. The larval
stage of the members of Chironominae required 16-
29 days and 20-40 days in summer and winter re-
spectively, whereas tanypodid species needed 45-50
days and 55-72 days respectively. Oliver (1971)
stated that changes in the duration of the larval
period occur over a wide range of latitudes. With
increasing latitude, the duration of the larval period
became longer. Temperature was one of the main
factors responsible for this increase. According to
Jonasson (1965), Chironomus anthracinus Zetter-
stedt, was recorded to have a two year life cycle,
although some emerged after one year and growth
was reduced both in summer (related to low oxygen
concentration) and in winter (related to low
temperature). The 2-year life cycle in other species
was also reported by Hamilton (1965) and Saether
(1968). Butler (1982) reported a 7-year life cycle for
two Chironomus species in arctic Alaskan tundra
ponds. The processes involved in slowing down or
cessation of growth and development have not re-
ceived due attention. Developmental arrests with a
cessation of feeding in both summer and winter
were found by Hilsenhoff (1966), Jonasson & Kris-
tiansen (1967) & Armitage (1970). From these stu-
dies, it became evident that temperature itself was
not the factor which controlled larval life, it rather
played a key role in determining the length of larval
life, but other factors, particularly the availability of
food, were also important (Jonasson 1965; Forsyth
19720)
The pupal stage had not been seriously investi-
gated except for its taxonomic importance. As com-
pared to the larval stage, the duration of the pupal
stage is very brief, ranging from a few hours to a few
days (Forsyth 1971, Oliver 1971, Chaudhuri et al,
1983, Chaudhuri & Ghosh 1986, Chattopadhyay et
al. 1988) and temperature dependent (Mundie
1956; Hilsenhoff 1966), which is also confirmed by
this investigation (table 2). The pupal life varied
between 21-48 hours and 26-56 hours in Chirono-
175
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
minae and between 46-66 and 51-75 hours in Tany-
podinae in summer and winter respectively.
The life span of adult chironomids lasts for a few
days (Table 2), supporting Forsyth (1971) and Hein
& Schmulbach (1972). The duration of adult life of
one week (Mundie 1956) or several weeks (Oliver
1971) is more unusual.
The emergence period is related to the duration
of the larval period (Chaudhuri et al. 1983). The
adults are able to fly immediately after emergence
as shown by Mundie (1956), Hilsenhoff (1956),
Brundin (1966), Forsyth (1971) and Hein &
Schmulbach (1972). The investigations revealed
that the rate of emergence was highest during the
middle of the total emergence period conforming
the findings of Chaudhuri et al. (1983) & Chatto-
padhyay et al. (1988). According to Ali (1980), the
periodicity is caused by a complex interaction of
various environmental stimuli and endogenous
rhythms. The phenomenon appears to be typically
regulated and maintained by an internal clock (the
endogenous rhythms) which is said to be influenced
by external stimuli or exogenous factors (Corbet
1964). Temperature is regarded as the more impor-
tant factor determining diel emergence periodici-
ties in high arctic chironomids (Oliver 1968; Danks
& Oliver 1972). Similar observations are reported
for mosquitoes by Corbet (1966). In contrast, inten-
sity of light appears as the factor controlling the diel
emergence pattern of chironomids in the temperate
zones. This study revealed that Chironomus sa-
moensis emerged at dusk in summer (see also Mor-
gan & Weddell 1961), while a few emerged at dawn.
Dicrotendipes pelochloris and Sticotochirono-
mus obscurus dis not show a distinct periodicity in
emergence, occurring mostly throughout the day
(Singh and Harrison 1982). A bimodal pattern was
also found by Ali & Mulla (1979) & Ali (1980).
However, a deviation was noticed in the peak of
emergence related to the earlier onset of sunset in
Clinotanypus fuscosignatus as stated above. Light
intensity may probably be the inducing agent in the
emergence of D. pelochloris and S. obscurus as
shown by Oliver (1968). Since these two environ-
mental clues were immediately linked together,
their combined effects require further investiga-
tion.
There are several reports on sex-ratios deviating
from the normal 1 : 1. According to Palmén (1962)
and Lindeberg (1971) males emerge in lower
numbers than females. This investigation revealed
a higher percentage of females in the case of Dicrot-
endipes pelochloris, Clinotanypus fuscosignatus
and Procladius noctivagus reared from egg masses
(Table 3), as also was notified by Palmén (1962) and
by Singh & Harrison (1982), but the reverse situa-
tion with a preponderance of males occurred in
Chironomus samoensis and Stictochironomus obs-
curus, as was also found by Hein & Schmulbach
174
(1972), Chaudhuri et al. (1983) and Chattopadhyay
et al. (1988). A rhythm in emergence starting with
an excess of males and ending with female predom-
inance was recorded in most of the chironomids in
question, as shown by Miall & Hammond (1900)
and Downes (1969), but no such rhythm was ob-
served in Stzcotochironomus obscurus in the pres-
ent study.
ACKNOWLEDGEMENTS
We are grateful to Prof. Bruno Rossaro, Depart-
ment of Zoology, University of Milan (Italy) for
kindly going through the manuscript and to Drs. B.
K. Tikadar, Director of Zoological Survey of India,
Calcutta, P. S. Cranston of British Museum (Natural
History), London (at present ANIC, CSIRO, Can-
berra, Australia) and D. Colless of the later Institute
for providing types of the chironomids for compar-
ison. Sincere thanks are due to the Council of Scien-
tific and Industrial Research, Government of India,
for financial assistance. We also thank Prof. Ole A.
Saether, University of Bergen, Norway for review-
ing the manuscript and his valuable suggestions.
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Saether, O. A., 1977. Taxonomic studies in Chironomidae;
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Saether, O. A., 1980. Glossary of chironomid morphology
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Sasa, M., 1978. A comparative study of adults and imma-
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Sasa, M., 1979. A morphological study of adults and im-
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148.
1727
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Sasa, M., 1983. Studies on chironomid midges of the Tama
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Sasa, M., 1985. Studies on Chironomid Midges of some
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Chironomidae). - Research report of the National
Institute of Environmental Studies, Japan 83: 25-99.
Sasa, M. & H. Hasegawa, 1983. Chironomid midges of the
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Sasa, M. & M. Kikuchi, 1986. Notes on the chironomid
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Sasa, M. & J. E. Sublette, 1980. Synonymy, distribution and
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178
Thienemann, A., 1954. Chironomus Leben, Verbreitung
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Received: 22 September 1989.
Revised version accepted: 19 July 1990.
CHAUDHURI & CHATTOPADHYAY: Chironomids
Figs. 1-5. Adults of Cryptochironomus judicius sp. n. 1, Frontal tubercles; 2, wing; 3, fore tibial scale; 4, tergites I-VI,
5, male hypopygium.
179)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
=
N
ER
ZA
ES EL
Figs. 6-11. Pupa of Cryptochironomus judicius sp. n. 6, Frontal apotome; 7, antennal sheath of male; 8, thoracic horn;
9, abdomen; 10, tergite II; 11, anal fin and genital sac of male.
180
CHAUDHURI & CHATTOPADHYAY: Chironomids
Figs. 12-21. Larva of Cryptochironomus judicius sp. n. 12, Head capsule; 13, antenna; 14, S I; 15, premandible; 16,
mandible; 17, prementohypopharyngeal complex; 18, mentum; 19, ventromental plate; 20, posterior abdominal seg-
ments; 21, claws of posterior parapods.
181
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 22-25. Adult of Harnischia tenuitubercula sp. n. 22, Frontal tubercles; 23, wing; 24, fore tibial scale; 25, male
hypopygium.
182
CHAUDHURI & CHATTOPADHYAY: Chironomids
Figs. 26-28. Adult of Paracladopelma aratra sp. n. 26, Wing; 27, fore tibial scale; 28, male hypopygium.
185
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 29-32. Adult of Paracladopelma sacculifera sp. n. 29, Frontal tubercles; 30, wing; 31 fore tibial scale; 32, male
hypopygium.
184
CHAUDHURI & CHATTOPADHYAY: Chironomids
EP,
Figs. 33-36. Adult of Polypedilum circulum sp. n. 33, Wing; 34, fore tibial scale; 35, abdomen; 36, male hypopygium.
185
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 37-40. Adult of Stenochironomus longipalpis (Kieffer). 37, Wing; 38, fore tibial scale; 39, abdomen; 40, male
hypopygium.
186
CHAUDHURI & CHATTOPADHYAY: Chironomids
No
/
Figs. 41-46. Adult of Stictochironomus obscurus (Guha & Chaudhuri) comb.n. 41, Wing; 42, fore tibial scale; 43, male
hypopygium; 44, female antenna; 15, female genitalia; 46, dorsomesal, ventrolateral and apodeme lobes of gonapophysis
VII.
187
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
È Sa =
{a Sn
„allan,
Ya UN
zu >
RP
G
Su
)
< I, dj
m)
a
Figs. 47-55. Pupa of Stictochironomus obscurus (Guha & Chaudhuri) comb.n. 47, Frontal apotome; 48, antennal sheath
of male; 49, antennal sheath of female; 50, thoracic horn; 51, abdomen; 52, tergite II; 53 anal fin and genital sac of male;
54, anal fin and genital sac of female; 55, caudolateral spur of segment VIII.
188
CHAUDHURI & CHATTOPADHYAY: Chironomids
> AW „ul = 7
64
Figs. 56-66. Larva of Stictochironomus obscurus (Guha & Chaudhuri) comb.n. 56, Head capsule; 57, antenna; 58, S I;
59, pecten epipharyngis; 50, premandible; 61, mandible; 62, prementohypopharyngeal complex; 63, mentum; 64,
ventromental plate; 65, posterior abdominal segments; 66, claws of posterior parapods.
189
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 67-73. Adult of Clinotanypus fuscosignatus (Kieffer). 67, Wing; 68, fore tibial spur; 69, mid tibial spur; 70,
abdomen; 71, male hypopygium; 72, gonostylus of male; hypopygium; 73, female genitalia.
190
CHAUDHURI & CHATTOPADHYAY: Chironomids
VA,
MR
À.
Zi
Z
ZA
SS
Figs. 74-80. Pupa of Clinotanypus fuscosignatus (Kieffer). 74, Frontal apotome; 75, antennal sheath of male; 76,
antennal sheath of female; 77, thoracic horn; 78, abdomen; 79, anal fin and genital sec of male; 80, anal fin and genital
sac of female.
WO
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 81-90. Larva of Clinotanypus fuscosignatus (Kieffer). 81, Head capsule; 82, antenna; 83, apex of antenna; 84,
mandible; 85, maxilla, 86, apex of maxilla; 87, ligula and paraligula; 88, mentum and M appendage; 89, posterior
abdominal segments; 90, claws of posterior parapods.
192
CHAUDHURI & CHATTOPADHYAY: Chironomids
Figs. 91-97. Adult of Procladius noctivagus (Kieffer). 91, Wing; 92, fore tibial spur; 93, mid tibial spur; 94, hind tibial
comb; 95, abdomen; 96, male hypopygium; 97, female genitalia.
195
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 98-104. Pupa of Procladius noctivagus (Kieffer). 98, Frontal apotome; 99, antennal sheath of male; 100, antennal
sheath of female; 101, thoracic horn; 102, abdomen; 103, anal fin and genital sac in male; 104, anal fin and genital sac
in female.
194
CHAUDHURI & CHATTOPADHYAY: Chironomids
Figs. 105-112. Larva of Procladius noctivagus (Kieffer). 105, Head capsule; 106, antenna; 107, mandible; 108, maxilla;
109, ligula and paraligula; 110, mentum and M appendage; 111, posterior abdominal segments; 112, claws of posterior
parapods.
195
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
196
HENK vAN MASTRIGT
Wamena, Indonesia
NEW (SUB)SPECIES OF DELIAS FROM THE
CENTRAL MOUNTAIN RANGE OF IRIAN JAYA
(EEPID OPMER A EIB REID ANE)
Mastrigt, H. J. G. van, 1990. New (sub)species of Deltas from the central mountain range
of Irian Jaya (Lepidoptera: Pieridae). - Tijdschrift voor Entomologie 133: 197-204, figs. 1-
17. [ISSN 0040-7496]. Published 14 December 1990.
Delias sinak sp. n., D. sigit sp. n., D. bobaga sp. n. and D. catocausta eefi subsp. n. are
described from the western part of the central mountain ranges in Irian Jaya, Indonesia. The
recent acquisition of new Delias material from Abmisibil in the Star Mountains of Irian Jaya,
leads to the recognition of the following new taxa: D. oktanglap sp. n., D. wollastoni
abmistbilensis subsp. n., D. luctuosa kuning subsp. n. and D. nieuwenhuisi poponga subsp.
n. Deltas nieuwenhuisi nom. nov., a replacement name for D. leucias roepkei Nieuwenhuis
& Howart (nec Benneth, 1956), is raised to specific rank.
Henk van Mastrigt, Kotak pos 25, Wamena 99501, Indonesia
Keywords. - Lepidoptera; Pieridae; Delias; new species; Irian Jaya; Indonesia.
From the beginning of this century quite a few
expeditions succeeded to enter the interior of Irian
Jaya (former Dutch New Guinea). The butterflies
of the genus Delias collected during these expedi-
tions provided material for the description of new
species from the Arfak Mountains, Weyland Moun-
tains, Wissel Lakes, the surroundings of the Car-
stensz Peak, the Baliem Valley, Mt. Goliath and the
Star Mountains.
The Star Mountains in Irian Jaya are one of the
central mountain ranges of which no reports on
Deltas have been published. The results of the
Dutch expedition to the Star Mountains in 1959
were poor and never published, as far as Delias is
concerned. In fact, most information about Delias
from the Star Mountains concerns the Papua New
Guinea part of this range, where the late Ray Straat-
man and others collected at Telefomin, close to the
Indonesian border. Results from the Papua New
Guinea part of the Star Mountains are found in
RMNH, BMNH and several private collections.
Since 1978 I have collected in the Star Mountains
of Irian Jaya (fig. 17). The weather in these moun-
tains is often cloudy and rainy, so that only small
numbers of Deltas can be collected in a short time,
contrary to more western ranges of the central
mountains where the weather is usually sunnier.
Besides that, the mountains are rough and steep,
which does not make travelling easy.
Further exploration of the extensive central
mountain range of Irian Jaya, with its isolated val-
leys led to the discovery of new taxa.
In this paper three new species and one new
subspecies from the western part of the central
mountain ranges are described. My recent acquisi-
tions of Delias from the Irian Jaya Star Mountains
and study of additional museum material led to the
recognition of one new species and three new sub-
species from this area. Finally, a previously des-
cribed subspecies is raised to specific rank.
Depositories
The abbreviations given below have been used in
the list of material and throughout the text.
BT - Private collection of Bernard Turlin, Andrésy,
France; CT - Private collection of Collin Treada-
way, Limbach, BRD.; DM - Private collection of
David Mannering, Staplehurst, Kent, U.K.; DR —
Private collection of David Rushen, E. Sussex, U.K.;
ERII - Private collection II of Eduard Reissinger,
Kaufbeuren, BRD.; GG - Private collection of Fred
Gerrits, Brisbane, Australia; HM - Private collec-
tion of the author, Jayapura, Indonesia; MZB -
Museum Zoologicum Bogoriense, Bogor, Indone-
sia.; RG - Private collection of Robert Gotts, Sid-
ney, Australia.; RMNH - Nationaal Natuurhisto-
risch Museum (Rijksmuseum van Natuurlijke
Historie), Leiden, The Netherlands; ZMA - Insti-
tuut voor Taxonomische Zoölogie (Zoölogisch Mu-
seum), Amsterdam, The Netherlands.
197
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 1-4. New (sub)species of the genus Delias; uppersides of male holotypes. 1, D. simak; 2, D. sigit; 3, D. bobaga;
4, D. catocausta eeft.
Figs. 5-8. New (sub)species of the genus Delias, undersides of male holotypes. 5, D. sinak; 6, D. sigit; 7, D. bobaga;
8, D. catocausta eeft.
198
VAN MASTRIGT: New Delias from Irian Jaya
Figs. 9-12. New (sub)species of the genus Delias, uppersides of male holotypes. 9, D. oktanglap; 10, D. wollastoni
abmisibilensis; 11, D.luctuosa kuning; 12, D. nieuwenhuisi poponga.
Figs. 13-16. New (sub)species of the genus Delias; undersides of male holotypes. 13, D. oktanglap; 14, D. wollastoni
abmisibilensis; 15, D.luctuosa kuning; 16, D. nieuwenhuisi poponga.
199
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
DESCRIPTIONS
Delias sinak sp. n.
Ges 1D 17)
Type material. - Holotype @: ‘Irian Jaya, route Mulia —
Sinak, vii.1989; don. Fr. Gerrits’, ZMA. - Paratypes: same
data as holotype, 2 6, GG; same data, but vii.1990, 6 3,
GG; 4 6, HM; Mulia, ix. 1990, 7 6, GG; 3 6 MZB.
Diagnosis
This species belongs to a subgroup of rather
small Delias, consisting of Deltas sagessa Fruh-
storfer, 1910 from East Papua New Guinea (includ-
ing its subspecies straatmani Schröder, 1977 from
Telefomin, Star Mountains and anjae Schröder,
1977 from the Arfak Mountains) and Deltas abro-
phora Roepke, 1955 from the Paniai-district. These
species are characterized by the black border of the
hind wing underside, bearing five rounded red
spots. The new species differs from these by the
absence of the red spot at the base of the yellow (and
white) discal band in the hind wing underside.
Description
Male. - Upperside fore wing white. Black border,
with regularly curved inner edge, from discal cell at
vein M, to margin border at 2A. Upperside of hind
wing translucent white with narrow (1.5 mm) black
border. Underside of fore wing black with a white
area (some grey-black diffusion), from middle of
tornus to vein CU,, sometimes entering discal cell,
three subapical yellow spots followed by two mar-
ginal ones. Underside of hind wing black, 5 mm
wide, yellow discal band from middle of tornus to
vein 2A, becoming narrower from vein M,, not
reaching anal border. Terminal border with five
large, more or less round, red spots, slightly variable
in size and shape.
Length of fore wing: 22 mm.
Female unknown.
Derivation of name. - ‘Sinak’: a noun in apposi-
tion. This butterfly occurs along the route from
Mulia to Sinak and received its name from the latter
locality.
Delias sigit sp. n.
(igs) 25, Oxley)
Type material. - Holotype @: ‘Irian Jaya/Centr. Berg-
land/Ilaga, River Jila/ 7.vi.1988/Henk van Mastrigt’
ZMA. - Paratypes: same data as holotype, 2 6, HM; same
data but 9.iv.1988, 5 g, ZMA; 6 4, MZB; 22.v.1988, 4 4,
HM; 2 6, BT; 27.viii.1988, 2 4, MZB; 2 6, HM; 1 6, BT;
2.v.1990, 2 6, HM; 4.v.1990, 3 6, HM; Ilaga, River Mum,
22.1.1989, 1 8, ZMA; Ilaga, 10.viii. 1988, 1 6, MZB; 1 4,
HM; 1 6, BT; 1 4, GG; Mulia-Sinak, vi-vii.1989, 2 8, GG;
Beoga, x11.1989, 7 8, GG; 1 6, HM; Mulia, ix. 1990, 10 6,
GG.
200
Diagnosis
This species is similar to Delias mariae Joicey &
Talbot, 1916 and D. bobaga because of its its bright
and broad distal band on the hind wing underside,
which is white with large orange patches. It differs
from D. bobaga in lacking the large number of
yellow scales in cells 2A and 3A of hind wing un-
derside. It differs from D. mariae in having a yellow
basal streak, which is connected to the white costal
border. The basal streak of D. mariae sometimes
with a small whitish edge, but the costa is black
instead of white.
Description
Male. - Upperside of fore wing milky white with
broad black costal border, entering discal cell, black
LDC. Black border along termen, much broader at
apex than at tornus and has slightly serrate inner
edge, often with 1 or 2 subapical spots; second one
very small. Upperside of hind wing milky white and
translucent, narrow black border; whole pattern of
underside visible from above. Underside of fore
wing milky white with black border narrower than
on upperside, three yellow subapical and one or two
very small terminal spots. Large anterior part of
discal cell black, passing LDC. Base of wing with
some black and some blackish diffusion. Underside
of hind wing yellow to yellowish orange with white
veins, rounded black submarginal spot reaching
LDC. Broad black anal part with some greyish green
diffusion, connected to white costa, cutting off a
yellow basal streak. Black border from SC+R, to 2A
increasing in width from less than 1 mm to 3 mm.
Length of fore wing: 25-29 mm.
Female unknown.
Derivation of name. — ‘Sigit’, a noun in apposi-
tion. ‘Sigit-sigit’ is the vernacular word for but-
terfly, used by the Dani people in the Ilaga valley,
where this butterfly occurs.
Delias bobaga sp. n.
(figs. 3, 7, 17)
Type material. - Holotype @: ‘Kamu Valley, 1700 m,
ix.1989, don. Henk van Mastrigt, ZMA. — Paratypes:
same data, 1 g, MZB; 1 8, GG; 1 6, DM; Kamu Valley:
Ekemanida, River Tuka & Enau, 1700-1800 m, 23-
30.x.1990, 3 &, HM; River Mouna, 1.11.1990, 5 À, GG;
River Epugewo, 2.11.1990, 1 6, GG; Idadagi, River Migu,
21.11.1990, 1 @, HM; Idadagi, River Bedonaka, 20.iv.1990,
1 4, HM; River Budau, 10.xi.1989, 1 4, GG; River Ode,
2.11.1990, 3 8, GG; Mapia area: River Okagou, 7.xii. 1989,
1 8, GG; 17.xii.1989, 1 4, HM; River Otika, 8.xii. 1989,
1 6, HM; River Yatuwou, 9.xii.1989, 1 6, GG; River
Gudopena, 11.xii.1989, 1 6, GG; River Ude, 13.xii.1989,
1 6, GG; River Piyakoywa, 1 8, GG; River Okagou &
Yatuwou, 25-26.xi.1989, 1 6, HM; Mapia, River lyage,
13.xii. 1989, 1 6, HM.
Diagnosis
This species is closely related to D. sigit sp. n.,
Delias mariae Joicey & Talbot, 1916 (including its
subspecies D. mariae boschmai Roepke, 1955) and
D. mariae menooensis Joicey & Talbot, 1922, with
which it occurs together in the Kamu Valley. How-
ever, it differs from those in the large number of
yellow scales in the cells 2A and 3A and in a much
larger basal spot on the underside of the hind wing.
Description
Male. - Upperside of fore wing white to creamy
white with broad, black costal border, entering dis-
cal cell, black LDC. Black border along termen,
much broader at apex than at tornus and has a
strongly serrate inner edge, sometimes absorbing
black LDC. Upperside of hind wing creamy white
and translucent with narrow black border; whole
pattern of underside visible from above. Underside
of fore wing milky white with black border, more
reduced than on upperside, three red subapical
spots and a red terminal one. Anterior part of discal
cell black, via M, connected to black border, separ-
ating a small, yellowish spot. Underside of hind
wing yellow with white veins and black submargi-
nal spot just outside discal cell. From middle of costa
a broad, curved, black line extends through inner
part of discal cell along yellow green anal area to the
bottom of cell CU, where it is connected by a thin
black line to the broad, black border, which runs
from RS to 3A. Inner part of costa white. Base
yellow with small black triangle at inner part.
Length of fore wing: 27-29 mm.
Female unknown.
Derivation of name. - ‘Bobaga’, a noun in appo-
sition, is the vernacular word for butterfly used by
the Ekagi people in the Paniai district, where this
butterfly occurs.
Delias catocausta eefi subsp. n.
(figs. 4, 8, 17)
Type material. - Holotype @: Irian Jaya/Tembagapura
2000 m/17.xi. 1979/Van Mastrigt’, ZMA. - Paratypes:
same data, 1 8, DR; same data but 17.xi.1979, 1 6, HM;
1 4, MZB; Tembagapura, 2000 m, 1980, 2 6, RG; 1983,
1 8, GG; Tembagapura, River Wanogong, 2000 m, 1989,
16, HM.
Diagnosis
Apart from the nominate form D. catocausta
catocausta Jordan, 1912, only D. catocausta niger-
rima Roepke, 1955 is known, from the Andrea
River, just 50 km west of Mt. Goliath (c. 140° E.).
The newly described subspecies is also from the
southern side of the central mountain range of Irian
Jaya, but occurs much more to the west: on the south
sides of the Puncak Jaya (Carstensz Peak), c. 137°
VAN MASTRIGT: New Delias from Irian Jaya
E. It differs from the two above mentioned ones in
the grey (instead of brown) ground colour on the
underside of fore and hind wing, in the red (instead
of yellow) colour of the basal spot on hind wing
underside and in having white veins.
Description
Male. — Upperside of fore wing milky white,
black tornus, thick, black LDC and black border
along termen; this black border is much broader at
apex than at tornus, has serrate inner edge and
bears 2 poorly developed subapical spots. Basis of
fore wing with some blackish diffusion. Upperside
of hind wing milky white with very narrow black
border. Underside of fore wing grey at innerside;
the grey colour is connected by black veins to black
border along termen; black border with three red
subapical spots and two very small terminal ones.
Inner edge of border serrate. A white band with
some grey diffusion separates dark inner part from
black border, widening from less than 1 mm on top
to 7 mm along tornus. Underside of hind wing grey
with white veins and small black spot close to discal
cell on vein M,. Cell along costa, inner part of discal
cell and cells 2A and 3A form a black hook with
small red basal spot and greyish green diffusion
along anal area in 3A. Narrow black border from
cell RS to anal area is undulate at inner edge and
separated from grey discal part by white line in cell
RS, getting broader in M,, M,, CU, and somewhat
narrower again in CU. Border between this white
line and grey innerpart vague and diffuse from M,
LCOS,
Length of fore wing: 27 mm.
Female unknown.
Derivation of name. - This subspecies is named
in honour of my father, the late Mr. Eef van Mas-
trigt, who awakened my interest in nature and es-
pecially in butterflies.
Delias oktanglap sp. n.
(figs. 9, 13)
Type material. - Holotype @: ‘Irian Jaya/Sterren Ge-
bergte/ Abmisibil 1920 m/ River Oktanglap/27.ii.1989/-
Henk van Mastrigt’, ZMA. - Paratypes: same data, 3 @,
ZMA; same data, but 11.ix.1985, 4 &, HM; 12.ix.1985, 1
6, HM; 14-16.ix.1985, 6 @, ZMA; 2 6, HM; 1 6, ZMA;
6.v1.1986, 4 4, RMNH; 29.iv.1986, 1 4, HM; 2.v.1986, 1
3, HM; 22.x.1986, 4 &, RMNH; 2 3, HM; 17.vii.1987, 3
6, HM; 11.iii.1988, 2 4, MZB; 16-19.iii.1988, 7 6, ZMA;
3 d, BT; 20.11.1988, 2 6, HM; Batimban, River Okkim,
1850 m, 14-16.11.1987, Van Mastrigt, 6 @, RMNH; same
data, but ix.1987, 2 @, ZMA; 19-21.iii.1988, 2 &, ZMA; 13-
15.iv.1988, 1 4, BT; v.1988, 2 6, MZB; 19-20.viii.1990, 2
8, HM; Abmisibil, River Takpalngi, 17-25.ix.1988, 2 3,
MZB; 1 8, BT; Abmisibil, River Okpeti, 1800 m,
14.ix.1986, 1 g, HM.
201
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Diagnosis
This new species belongs to the cuningputi
group, consisting of D. cuningputi (Ribbe, 1900),
D. konokono Orr & Sibatani, 1986, D. chimbu Orr
& Sibatani, 1986, D. fascelis fascelis Jordan, 1912, D.
fascelis ibelana Roepke, 1955, D. citrona Joicey &
Talbot, 1922 and D. jordanı Kenrick, 1909, as de-
fined by Orr and Sibatani (1986). It is distinguished
from all those by its very reduced black border on
the fore wing upperside and from cuningputi and
konokono by its continuing submarginal light band
on the hind wing underside.
Description
Male. - Upperside of fore wing white, black
border from 1.5 mm outside discal cell to tornus,
inner edge straight and slightly serrate. Very thin
bar at top of discal cell. Upperside of hind wing
white with black border from SC+R, to CU,, 3 mm
wide in M,, narrowing to both ends. Underside of
fore wing white in bottom 1/3, with black diffusion.
Large black border connected with nearly totally
black discal cell, enclosing three subapical yellow
spots and three yellow to white marginal spots —
with thin white line to margin border. Veins be-
tween discal cell and border partly black. Underside
of hind wing black, divided by an irregular, 2-6 mm
wide, curved white discal band, broken in M,. Ter-
minal border white with a yellow basal streak. Inner
part of hindwing black with two yellow spots and
some yellowish diffusion. On white discal band one
yellow spot just above black connection, five yellow
streaks in underpart. Black border with undulate
edge at innerside, having a row of six yellow spots
in a line with the above mentioned yellow spot and
streaks on white band.
Length of fore wing: (25) 28 — 29 mm.
Female unknown.
Derivation of name. - ‘Oktanglap’ (a noun in
apposition) is the name of the creek from where the
holotype originates.
Delias wollastoni abmisibilensis subsp. n.
(figs. 10 & 14)
Type material. - Holotype &: ‘Irian Jaya/Abmisibil
1920 m/River Oktanglap/ 11.1x.1985/Van Mastrigt’,
ZMA. - Paratypes: same data as holotype, 1 8, HM; same
data, but 1880-1920 m, 15.ix.1985, 1 4, HM; 22.x.1986, 1
3, MZB; 1 4, HM; 17-19. vii.1987, 1 6, ZMA.
Diagnosis
Adding this new subspecies from the Star Moun-
tains in Irian Jaya, three separated populations of
Delias wollastoni Rothschild, 1915 are now known,
the other two being: D. wollastoni wollastoni from
the southside of the Puncak Jaya (Carstensz Peak)
(1 male in BMNH) and Beoga (1 male in GG) and
D. wollastoni bryophila Roepke, 1955 from the
202
surrounding of Mt. Trikora (Mt. Wilhelmina)
(good series of males in RMNH and ZMA and two
males in HM). The new subspecies differs from the
other two subspecies by its golden yellow colour of
the fore wing underside and more vivid coloration
of hind wing underside.
Description
Male. - Upperside of fore wing white, greyish
black veins and some grey diffusion. Costal border
black. Black border from top of discal cell to tornus
encloses black LDC, two white subapical spots, two
or three small terminal ones. Border reduced in cells
M,, CU, and CU, and slightly entering along dor-
sum. Upperside hind wing white with grey veins
and some grey diffusion, particularly at inner edge
of broad black border in cell M, and M,. Underside
of fore wing golden yellow, black border much nar-
rower than on upperside, regularly curved at inner
edge, encloses three large yellow subapical spots
and five smaller yellow terminal ones. Costal border
black; at top of LDC large black triangle which is
narrowly connected to black border. Underside of
hind wing with base and basal half of costa grey,
bordered by SC+R,. Large basal maroon coloured
patch below SC+R,, oblong patch on LDC and
bowed, undulate postmedian band of same colour.
Postmedian band consists of intervernal patches
from RS to CU,. Anal area densely covered with
golden yellow scales; grey becoming pinkish grey in
outer third of discal cell and in adjacent cells. A
strong and rather straight median band extends
from costa to 1A+2A, including the maroon brown
streak on LDC. Space between this median and
postmedian band filled with light pinkish grey, den-
sely covered with dark grey-brown diffusion. Sub-
terminal area light pinkish grey with darkened ter-
men, more or less dusted with golden-yellow scales
mainly on tips of veins.
Length of fore wing: 29 mm.
Female unknown.
Derivation of name. - Abmisibil is the name of
the village in the Star Mountains, which is close to
the sites where this butterfly occurs.
Delias luctuosa kuning subsp. n.
(figs. 11 & 15)
Type material. - Holotype @: ‘Irian Jaya/Star Moun-
tains/ Abmisibil 1800 m/River Oktero/ 11.1x.1985’, ZMA.
— Paratypes: Abmisibil, River Okmi, 31.iii.1985, Sam Mo-
nareh; 1 6, HM; Abmisibil, River Oktanglap, H. van
Mastrigt, 1 @, HM; Langda, River Bibwe, 1800 m,
6.iv.1989, H. van Mastrigt, 1 d, HM; same data, but 17-
19.iv.1989, 1 6, HM.
Diagnosis
Three populations of D. lwctwosa Jordan, 1912
have been named previously, viz. D. /uctuosa arch-
boldi Roepke, 1955 from the Baliem Valley (also
from the more western Ilaga Valley), D. lwctuosa
luctuosa from Mount Groome, Papua New Guinea,
and D. luctuosa mizukamit Okana, 1989 from
Simbu Province, Papua New Guinea. With the new
population from Langda and Abmisibil a bridge is
made between the population from central Irian
Jaya and the two populations from Papua New
Guinea. The new subspecies differs from the three
above mentioned in the golden-yellow colour of the
male fore wing underside.
Description
Male. - Upperside of fore wing white with black
costal border, slightly entering discal cell. Terminal
border very broad in cells M,, M, and M;, narrower
towards tornus, where it slightly curves along dor-
sum, provided with irregular inner edge, enclosing
two small white subapical spots. Veins visible, par-
ticularly those on underside of discal cell. Upperside
of hind wing translucent white with black border,
which is narrow in cell RS and M,, undulate at vein
M, and M; and broad in cells CU,, CU, and 2A.
Underside of fore wing deep yellow, turning to
white along dorsum, black border along inner edge
in cell 2A. Costal border black. Large LDC con-
nected to black border in cell M;, demarcating two
yellow spots with black diffusion in cells M, and M,.
Black border encloses three dark yellow subapical
spots and two or three small terminal spots. Under-
side of hind wing with white ground-colour. Black
costal border connected to broad black anal border,
including small yellow pear-shaped basal spot.
Postdiscal black band (width 3 mm) curving from
anal border to costa at vein SC+R, and entering
broadly cell RS; outer edge bordered with small red
un
+
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VAN MASTRIGT: New Delias from Irian Jaya
line. White discal part with pink diffusion, darker
at innerside; veins white. Discal border consists of
five black triangles at veins, touching red line and
demarcating six white, triangular to v-shaped, spots.
Length of fore wing: 27-29 mm.
Female unknown.
Derivation of name. — A noun in apposition.
‘Kuning’ is the Indonesian word for yellow and
refers to the yellow underside of the fore wing, the
characteristic feature of this butterfly.
Delias nieuwenhuisi nom. nov.
Delias leucias roepkei Nieuwenhuis & Howart, 1969: 86.
Holotype 4: Mandated New Guinea, Telefomin, 1700
m, 5.viii. 1962 (Natuurhistorisch Museum, Rotterdam)
(preoccupied by D. mira roepkei Benneth, 1956).
Discussion
Since the subspecific name roepkez in Delis leu-
clas roepkei Nieuwenhuis & Howarth is preoccu-
pied by Benneth’s name, the replacement name
meuwenhuist is proposed here, in honour of Mr. E.
J. Nieuwenhuis, who was the first to describe this
taxon. Further, since nzeuwenhuisi and leucias s. str.
are sympatric in the Star Mountains along river
Okkim at Abmisibil, Irian Jaya and at Telefomin,
Papua New Guinea, they cannot be regarded as
subspecies of one species, therefore the first one is
raised to specific rank: D. nieuwenhuisi.
The differences between the specimens from
Telefomin (types of nieuwenhuisi) and those from
Abmisibil lead to the recognition of two subspecies:
D. nieuwenhuisi nieuwenhuisi from Telefomin and
D. nieuwenhuisi poponga subsp. n. from Abmisibil
(see below).
Fig. 17. Distribution of
new (sub)species of the ge-
nus Delias. a, Kamu valley
(type locality of bobaga); b,
Tembagapura (type locality
of catocausta eeft); c, llaga
(type locality of sgit); d,
Sinak (type locality of
sinak); e. Langda (para-
types of /uctuosa kuning); f,
Abmisibil (type locality of
oktanglap, wollastoni ab-
misibilensis and luctuosa
kuning).
159)
03
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Delias nieuwenhuisi poponga subsp. n.
(figs. 12, 16)
Type material. - Holotype 8: ‘Irian Jaya/Sterren-
gebergte/Batimban/ca 1900 m/ River Okkim, v.1988/ H.
van Mastrigt’, ZMA. - Paratypes: same data, 9 8, HM; 1
é, ZMA; same data, but 24.xii.1985-7.1.1986, 3 6, HM;
same data, but 1-5.v.1986, 14 6; 2 6, DR; 6 g, RMNH;
same data, but 14-16.ii.1987, 6 @, ZMA; same data, but
ix.1987,7 6, HM; 1 8, BT; 6 g, ZMA; same data, but 19-
21.iii 1988, 14 À, HM; 6 @, BT; same data, but 13-
15.iv.1988, 2 6, FG; 1 6, HM; 1 4, CT; same data, but 22.v-
5.vi.1988, 2 6, HM; same data, but 11-13.viii.1988, 4 3,
HM; same data, but vi.1986, 2 6, ERII; same data, but 19-
20.viii.1990, 4 6, HM; 4 4, MZB; Takpalngi, 17-
25.ix.1988, H. van Mastrigt, 6 4, HM; Ferohir, 5.v.1986,
H. van Mastrigt, 1 @, ZMA.
Diagnosis
This subspecies differs from nominate nieuwen-
huisi in three characters. Upperside and underside
of fore wing having narrow black borders (even
narrower than in leucias); having upperside of fore
wing with a black vein, connecting border with
midcell and underside of hind wing with a large
posterior red strike and a thin anterior white line
in the basal spot.
Description
Male. - Upperside of fore wing white with nar-
row black border, 5 mm in cell R, to 1 mm in cell
CU. Inner edge slightly serrate, not curved from
cell R, to tornus. Black vein R, connects border with
discal cell, demarcating a small white spot. Apical
and marginal spots absent. Upperside of hind wing
white, 1-2 mm wide, black border. Underside of fore
wing yellow, narrow black border as broad as on
upperside, having four yellow apical spots. Ground-
colour orange yellow in cells R,, M,, M,, M;, and
CU,, lemon yellow in discal cell and pale yellow to
creamy white in cells CU, and 2A. Underside of
hind wing with large creamy discal area, covering
nearly whole discal cell and large inner parts of cells
RS, M,, M,, M; and CU,, just entering cell CU.
Discal area surrounded by black line, which is less
than 1 mm in cell RS, 2 mm in cell CU, and 3.5 mm
in discal cell. Red subbasal mark usually oblong,
sometimes more like an upside down ‘v. Al mm
wide red line runs from cell SC+R, to CU,, parallel
to and connected with black line which surrounds
discal area. Discal border consists of a, 1.5 mm wide,
white line, with at its outer edge a narrow black line
from cell M, to CU,, which is more than 1 mm in
cells M; and CU, only. Basal spot anteriorly white,
posteriorly red. Anal margin appearing dirty pale
yellow due to low number of black scales (compared
to other species of this subgroup). Concentration of
white scales along vein 3A divides yellow anal area.
Length of fore wing: 22-24 mm.
Female unknown.
Derivation of name. - A noun in apposition.
‘Poponga’ is the vernacular word for butterfly used
by the people in the western part of the Star Moun-
tains, where this butterfly occurs.
ACKNOWLEDGEMENTS
I gratefully acknowledge the help of Dr. Fred Gerrits,
Mr. David Mannering, Mr. Bernard Turlin and Mr. Collin
Treadaway, Mr. David Rushen, who allowed me to study
their private collections and the assistance of Dr. Eduard
Reissinger and Dr. Atuhiro Sibatani. Many thanks are
extended to the Instituut voor Taxonomische Zoölogie
(Zoölogisch Museum), Amsterdam, for the good and
helpful cooperation, especially to Mr. L. van der Laan for
the preparation of the photographs.
The cooperation with the Museum Zoologicum Bogo-
riense, Bogor and the Department of Forestry, Irian Jaya,
in the study of butterflies from Irian Jaya is gratefully
acknowledged.
REFERENCES
Bennett, N. H., 1956. A new Deltas from the central
highlands of New Guinea (Lepidoptera, Pieridae). —
The Entomologist 89: 81-83, pl. 3.
Fruhstorfer, H., 1910. Familie: Pieridae, Weisslinge. - In:
A. Seitz, Die Gross-Schmetterlinge der Erde 9: 123-
135, 183-190, (1927) 1108.
Joicey, J.J. & G. Talbot, 1916. New lepidoptera from Dutch
New Guinea. - Annals and Magazine of Natural His-
tory (series 8) 17: 68-72.
Joicey, J. J. & G. Talbot, 1922. New forms of the genus
Deltas (Pieridae) from New Guinea, Ceram and Buru.
— Bulletin of the Hill Museum 1 (3): 303-319.
Jordan, K., 1912. On new or little-known Delias from
New Guinea. - Novitates Zoologicae 18 (1911): 580-
DOSI
Nieuwenhuis, E. J. & T. G. Howart, 1969. On some but-
terflies from the Indo-Pacific region. - Entomolo-
gische Berichten, Amsterdam 29: 85-88.
Kenrick, G. H., 1909. Descriptions of some new species
of the genus Delias from North New Guinea, recently
collected by Mr. C. E. Pratt. - Annals and Magazine of
Natural History (series 8) 4: 176-183, pls 6, 7.
Okana, K., 1989. Descriptions of four new butterflies of
the genus Deltas (Lep.: Pieridae), with some notes on
Delias. - Tokurana 14 (3/4): 1-6.
Orr, A. G. & A. Sibatani, 1986. A revision of the Delias
aroae-cuningputi complex (Lepidoptera, Pieridae) 1.
The D. cuningputi group. - Tyô to Ga 37 (1): 1-14.
Ribbe, C., 1900. Neue Lepidopteren aus Neu-Guinea. —
Insektenbörse 17: 329-330.
Roepke, W. K. J., 1955. The butterflies of the genus Delias
Hübner (Lepidoptera) in Netherlands New Guinea. —
Nova Guinea, n. s. 6 (2): 185-260.
Rothschild, W., 1915. — In: W. Rothschild and J. H. Dur-
rant, Lepidoptera of the British Ornithologist's Union
and Wollaston Expeditions in the snow mountains,
southern Dutch New Guinea. - 168 pp. London, Ha-
zell, Watson & Viney, Ltd.
Schröder, H., 1977. Zwei neue Delias der sagessa-Gruppe
aus Neuguinea (Lep., Pieridae). - Entomologische
Zeitschrift Frankfurt am Mainz 87: 265-271.
Received: 18 July 1990
Accepted: 14 September 1990
204
ERIK J. vAN NIEUKERKEN
National Museum of Natural History, Leiden
THE TRIFURCULA SUBNITIDELLA GROUP
(LEPIDOPTERA: NEPTICULIDAE): TAXONOMY,
DISTRIBUTION AND BIOLOGY
Nieukerken, E. J. van, 1990. The Trifurcula subnitidella group (Lepidoptera: Nepticulidae):
taxonomy, distribution and biology. — Tijdschrift voor Entomologie 133: 205-238, figs. 1-
108, tab. 1. [ISSN 0040-7496]. Published 14 December 1990.
The nominal subgenus Trifurcula Zeller, 1848 is divided into two species groups: the
subnitidella group and the pallidella group. The subnitidella group is described and com-
prises T. subnitidella (Duponchel, 1843), and eight new species: austriaca (from Austria),
luteola (France), puplesisi (Soviet Union), coronillae (Spain), victoris (Spain), josefklime-
schi (Italy, southern Europe), iberica (Spain) and silviae (France). Males of all species and
females of six species are described and diagnosed, and for four species the larvae, mines
and biology are described as well: they make mines in stems of herbaceous or shrubby
Fabaceae (tribes Loteae, Coronilleae). The group is widespread in the western Palaearctic,
but most species occur in Southwest Europe. T. subnitidella is recorded for the first time
from 14 countries. The phylogeny of the group and subgenus are discussed, the pallidella
group is considered to be its sistergroup. A lectotype for T. orientella Klimesch, previously
confused with some species here described, is designated.
Dr. E. J. van Nieukerken, Rijksmuseum van Natuurlijke Historie, Postbus 9517, 2300 RA
Leiden, Netherlands.
Keywords. - Nepticulidae, stem-miners, taxonomy, phylogeny, Palaearctic, Fabaceae.
The genus Trifurcula Zeller, 1848 constitutes an
important part of the nepticulid fauna of the Me-
diterranean region with the total number of species
probably exceeding 60 (31 described species: van
Nieukerken 1986a). It is especially abundant in dry
open habitats, such as grasslands, maquis and gar-
rigue vegetations, where the larvae make leaf- or
stem-mines in shrubby and herbaceous plants, in
particular Lamiaceae (Labiatae) and Fabaceae (Le-
guminosae). A few species extend their ranges
northwards into northern and western Europe. Tri-
furcula remains the main nepticulid taxon in Eu-
rope which needs extensive taxonomic revision.
The genus was fully redescribed by van Nieu-
kerken (1986b), who recognized three subgenera:
Glaucolepis Braun, Levarchama Beirne and Trifur-
cula Zeller. The ten Northwest European species
have been treated by van Nieukerken & Johansson
(1990), but no comprehensive work for the remain-
ing European species is currently available.
The present paper is the first in a series of revi-
sions of Trifurcula, which follows a few papers
dealing with single species (van Nieukerken & Jo- .
hansson 1986, van Nieukerken 1987). The subge-
nus Trifurcula was revised by Klimesch (1953), who
recognized eight species, two of which are now
considered to be junior synonyms. Only two species
were added later, T. griseella Wolff, 1957 (later
recognized to be a junior synonym of T. subnitidella
(Duponchel, | 1843 ])) and 7. beirnei Puplesis, 1984,
resulting again in a number of eight described spe-
cies (van Nieukerken 1986a). These species only
represent a minority of the actual number, which
probably exceeds 25.
The described species are almost all relatively
uniformly coloured moths, without striking exter-
nal characters, and associated with various species
of brooms (Fabaceae, tribe Genisteae). T. subniti-
della forms an exception in having a conspicuous
external character — a brand of bright yellow scales
on the male forewing underside — and having an
association with herbaceous Fabaceae (Lotus: van
Nieukerken & Johansson 1990). In unidentified and
misidentified material from southern Europe and
western Asia eight species with similar external
features have been found, and following recent field
trips three of these species and T. subnitidella itself,
have been reared from larvae collected on plants
belonging to the tribes Loteae and Coronilleae. In
this paper these species are assembled into the T.
subnitidella species group, whereas the other pa-
laearctic species in Trifurcula s. str. (with the type
species T. pallidella (Duponchel, [1843])) are con-
sidered to form the T. pallidella species group.
205
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
When studying the material, representing the
new species below, several specimens were found to
be labelled as T. orzentella Klimesch, 1953. This led
me to re-examine T. orientella and select a lectotype
for this species, which clearly belongs to the pa//-
della group. Other specimens treated here were
found amongst material labelled as ‘T. immundella
(Zeller)’, ‘T. pallidella Zeller’, ‘T. serotinella Her-
rich-Schaffer’ and ‘Nepticula cryptella Frey’. Ap-
parently, previous workers rarely examined the un-
derside of the forewings of these tiny moths, which
exhibit such good characters in this group.
MATERIAL AND METHODS
Larvae were collected by cutting portions of the
stems of the hostplant, containing the mines. The
206
Figs. 1-6. Trifurcula spp., male habitus. - 1, T. austriaca, holotype; 2, T. luteola, paratype from type-locality; 3, T.
coronillae, holotype; 4, T. victoris, holotype; 5, T. subnitidella, Spain, Camino de Ojen; 6, T. josefklimeschi, holotype.
stems were put in plastic boxes with a layer of paper
tissue, in which the larvae could spin their cocoons.
Next the cocoons or full-grown larvae were trans-
ferred into glass jars with a layer of soil and some
moss. In the case of species from southern Europe,
these jars were kept indoors (in The Netherlands),
at room temperature, until emergence. Autumn col-
lected larvae of subnitidella were, after spinning
their cocoons, left in an outhouse for overwintering,
and were brought indoors in March.
Methods for preparation of the genitalia are lar-
gely the same as in van Nieukerken (1985). In this
group it is often very difficult to separate the aedea-
gus from the genitalia capsule and valvae, because
of the tight connection with membranes. The ae-
deagus has therefore only been severed in some
specimens, in order to study the complex morphol-
VAN NIEUKERKEN: Trifurcula subnitidella group
DEN
Wh Ws
Figs. 7-10. Trifurcula spp., males. - 7, T. silviae, habitus, holotype; 8, T. austriaca, underside, holotype; 9, T. subnitidella,
underside, Spain, Camino de Ojen; 10, 7. coronillae, underside, holotype. Andraconial patch arrowed in figs. 8-10.
ogy of aedeagal processes and cornuti; in some spec-
imens also the vesica was removed from the aedea-
gal tube to show the cornuti better. For identifica-
tion purposes, removing of the aedeagus is not
always necessary.
Larvae were treated with KOH 10% prior to
cleaning and mounting with Euparal on slides.
They were stained with Chlorazol black.
Line figures of genitalia were prepared with a
Zeiss Standard microscope or Zeiss Axioskop, with
camera and drawing apparatus, both from genitalia
in glycerin and permanent mounts. In most cases
cornuti are drawn separate from the aedeagus; only
the characteristic larger cornuti are depicted, the
smaller ones being almost similar in all species.
Line drawings of mines were made from herbarium
specimens, after boiling for some minutes in dis-
tilled water: frass may be better visible in such
soaked mines than in fresh ones.
Photographs of genitalia and larval head capsules
were taken with a Zeiss Axioskop microscope and
camera.
SEM micrographs were taken with a Jeol JSM
840A scanning electron microscope. Specimens
were air-dried, mounted on stubs and gold-coated.
Measurements of genitalia are taken with a Zeiss
Axioskop or Universal Microscope, at 200 X, and
are accurate at the nearest 5 um. All measurements
based on sample size of at least five specimens are
accompanied by mean, standard deviation and sam-
ple size in brackets.
Distribution maps are based on material exam-
ined, but for T. subnitidella also literature data and
personal communications have been used. Locality
names are spelled in accordance with the Times
Atlas of the World (Comprehensive edition, 1975
and later), deviating names on labels are cited in
brackets. For all localities the appropriate UTM
grid references (10 X 10 km, or 1 X 1 km for
authors’ records) are given. These are used for plot-
ting the distribution on the 50 X 50 km squares of
the maps provided by the ‘European Invertebrate
Survey’. The UTM grid references were obtained
directly from ‘Tactical Pilotage Charts 1:500,000',
topographical maps, or calculated from geographi-
cal coordinates (obtained from atlases, various
maps or US gazetteers), using the computer pro-
gram ‘UTM’ by Rasmont & André. Most of the
British records, however, are an approximation of
the correct UTM square, since they are based on
vice-county records and dots on a vice-county map,
provided by A. M. Emmet (in litt.).
Nomenclature of hostplants follows Heywood &
Ball (1968), but the family name Fabaceae is used
in stead of Leguminosae.
Abbreviations (codens) for depositories follow
Arnett & Samuelson (1986), with the addition of:
ETO (for collection E. Traugott-Olsen, Marbella,
207
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 11-16. Trifurcula subnitidella, male, details of wings, scanning micrographs. - 11, forewing underside with yellow
scale patch, some normal wing scales in upper right corner; 12, 13, details of androconial scales; 14, hindwing with frenulum
and group of ‘black’ scales; 15, idem, detail; 16, ultrastructure of ‘black’ scale. bs = black scales; ns = normal scale.
Spain), GBA (for collection G. Baldizzone, Asti,
Italy) and ZKVV (Zoologijos Katedra VPI, Vilnius,
Lithuania). Note that in the index of Arnett &
Samuelson the coden NHMW is misspelled as
NHMV. RMNH remains the coden for the Na-
tional Museum of Natural History, Leiden, Nether-
lands (formerly Rijksmuseum van Natuurlijke His-
torie). Genitalia slide numbers refer to the
numbering system of the author, unless accompan-
ied by another abbreviation.
208
TAXONOMIC PART
For descriptions of the genus Trifurcula Zeller,
1848 and its typical subgenus refer to van Nieu-
kerken (1986b) and van Nieukerken & Johansson
(1990). This subgenus is here divided into two spe-
cies groups, recognized according to the following
key:
1. Forewing underside in male with a patch of
yellowish androconial scales near base (except
in T. silviae); in addition hindwing costa often
with row or group of dark brown or black spe-
cial scales. Male genitalia: gnathos often atyp-
ical: either with serrations, keels, projections or
completely asymmetrical, occasionally simple
and symmetrical. Aedeagus usually with dorsal
lobe at right side; large cornuti (3) not curved
and horn-like (except in awstriaca). Larvae
stem-miners in tribes Loteae or Coronilleae
(Rabaccao) enn: subnitidella group
- Forewing underside and hindwing costa in
male without androconial or special scales.
Male genitalia: gnathos simple, symmetrical.
Aedeagus usually with dorsal lobe at left side;
at least one or two large cornuti strongly
curved, horn-like. Larvae stem-miners in
brooms: tribe Genisteae..... pallidella group
The following nominal species are here assigned
to the pallidella group: Trifurcula pallidella (Du-
ponchel, [1843]) (= incognitella Toll, 1936), T.
immundella (Zeller, 1839), T. serotinella Herrich-
Schäffer, 1855 (= confertella Fuchs, 1895), T. orien-
tella Klimesch, 1953, T. aurella Klimesch, 1953, T.
beirnet Puplesis, 1984 and T. squamatella Stainton,
1849 (= maxima Klimesch, 1953). Also the undes-
cribed species from Sumatra (see van Nieukerken
1986b) belongs here. This group will be the subject
of a future revision.
The Trifurcula subnitidella group
Description
Adult (figs. 1-7). - Forewing length 2-3.5 mm.
Antenna with 27-48 segments, in male with more
flagellar segments than in female. Forewing upper-
side uniformly pale or irrorate by dark tipped scales,
mixed with pale scales. No other colour pattern
present. Venation as described for subgenus (van
Nieukerken 1986b, van Nieukerken & Johansson
1990).
Underside of forewing in male with a distinct
patch of lamellar androconial scales near wing-base
(figs. 8-10), usually yellow or yellowish white, oc-
casionally with various tinges of yellow (this patch
is absent in T. silviae). The patch consists of scales,
which are ultrastructurally different from normal
wing scales (fig. 11) (see also van Nieukerken et al.
1990). In szbnitidella these scales have almost no
ultrastructure (figs. 12, 13), although remnants of
scale-ribs are visible on some scales. In coronillae
the three differently coloured areas also appear to
have different ultrastructure: the basal area is much
similar to the patch of szbnitidella (not illustrated),
but the scales in the central area have an irregular
pattern of ribs (fig. 19). Forewing costa often with
VAN NIEUKERKEN: Trifurcula subnitidella group
distinct costal fold on underside, in josefklimeschi
extremely large (figs. 20, 58).
Hindwing underside in male with terminal
velvet-like patch of raised scales (generic character)
(illustrated by van Nieukerken 1986b and van
Nieukerken et al., 1990). Costa of hindwing in sev-
eral species slightly enlarged (fig. 21), in some spe-
cies provided with a short or longer row of distinct
black androconial scales (figs. 57-60). These scales
also differ ultrastructurally from normal wing
scales (figs. 15, 16, 22). In fig. 15 both the black
scales and normal wing scales are shown, the ultra-
structure of the normal scale is illustrated in fig. 17.
The androconial scales are completely jet-black, or
dark-brown in sz/viae, and have spiny ribs, with the
intercalary area almost devoid of structure, apart
from some indistinct bulges (fig. 16). Male without
costal bristles, but often with long white hair-scales
instead.
Male genitalia (figs. 23-88 in part). - Vinculum
anteriorly rounded, truncate or slightly concave.
Uncus with central process pointed or terminally
widened, occasionally bilobed. Gnathos occurs in
two forms: either symmetrical, with rounded or
triangular central element, or asymmetrical and
complex, provided with keel(s) or with serrate mar-
gin or with a combination of both; anterior apo-
demes usually present, but not always well sclero-
tized. Valva elongate, more or less triangular, or in
some species with broader basal part. Aedeagus
joined by juxta (ventral process) to valvae and cap-
sule. Aedeagal tube asymmetric, complicate, taper-
ing posteriorly both in lateral and ventral view;
ventrally ending in a pointed or serrate (fringed)
medial carina, which hinges to the juxta, dorsally
ending ina pointed or spatulate tip, often indistinct,
and in addition usually with a dorsal lobe at right-
hand side, in austriaca at left-hand side. Vesica with
complex set of cornuti: a group of about 10-20 long
needle-like cornuti at right-hand side (dorsally in
lateral view), many smaller cornuti of variable
shape spread throughout vesica and usually three
larger cornuti near phallotrema: a long smooth
spine, often protruding from phallotrema, basally
connected to a smaller cone-shaped cornutus; in
addition another large, often irregular cornutus is
often hidden amongst the group of spine-like cor-
nuti, it may also protrude from the aedeagal tube.
Sometimes both large cornuti are very similar, and
other cornuti may be more reduced in size and
number. In coronillae the dorsal lobe has a spiny
process, which in normal slides is inseparable from
the cornuti (fig. 49).
Female genitalia (figs. 89-100). — In general as in
(sub)genus. Terminal segments either broad and
blunt, or slightly pointed. Ductus bursae relatively
wide; corpus long, covered with minute pectina-
tions; signa usually long and narrow: a thickened
margin encloses some rows of complete cells, out-
209
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 17-22. Trifurcula spp., male, details of wings. - 17, T. subnitidella, ultrastructure of normal wingscale from fig.
15; 18, T. coronillae, forewing underside with yellow patch, different areas numbered from wingbase; 19, idem,
ultrastructure of scale from area 2; 20, T. josefklimeschi, forewing underside with costal fold and androconial scale patch
underneath; 21, idem, hindwing with humeral lobe and ‘black’ scales, costal margin below; 22, idem, detail. bs = black
scales; f = fold; hl = humeral lobe; hs = hairscales.
side this margin there is a row of incomplete larger
cells. Ductus spermathecae with less than 5 coils.
Final instar larva (figs. 101-104). — Body ex-
tremely elongate, deep yellow in colour. Head with
frontoclypeus almost rectangular, tentorial arms
short. Labrum with two pairs of setae: a medial and
a lateral. Labial palpi three segmented. Antenna
with sensilla not cross-wise. Prothorax with small
ventral sclerite and a pair of narrow dorsal sclerites.
Thorax with respectively 13, 12 and 10 setae pairs,
210
abdominal segments 1-8 with 6, segment 9 with 3
and 10 with 2 setae pairs in all species examined.
Integument with reduced spinosity, usually only
spines ventrally and laterally upwards to the spir-
acles; spines usually short and inconspicuous. Anal
rods posteriorly bifid. The four species described
below can hardly be distinguished as larva, except
by size, length/width ratio of the headcapsule and
slight differences in spinosity.
Diagnosis
Males of the subnitidella group are recognized by
the presence of a yellow androconial patch on the
forewing underside, in some species in combination
with black scaling along the hindwing costa. In 7.
josefklimeschi the patch is hidden under a large
costal fold, in s#/vrae the patch is absent, but there
are rows of dark brown to black scales along the
margin of the forewing costal fold and along the
hindwing costa. Males of several species in the
subgenus Glaucolepis Braun (such as T. stoecha-
della (Klimesch) and T. saturejae (Parenti)) also
have patches of androconial scales on the underside
of the forewing: they are either farther away from
the wing base or extend over almost the whole wing
surface; these species also differ in venation and
genitalia (see van Nieukerken & Johansson, 1990).
Females cannot be differentiated externally from
other Trifurcula species. The female genitalia differ
from other Trifurcula s. 1. species by a combination
of the typical shape of signa as described above and
the relative short ductus spermathecae with few
coils. However, some species of the Trifurcula pal-
lidella group have similar female genitalia.
Biology
The biology of this group has not been described
prior to the recent discovery of the biology of T.
subnitidella (van Nieukerken & Johansson 1990).
The hostplant and larva of four of the species des-
cribed here are known through rearing. The larvae
of these species make stem-mines in herbaceous or
shrubby species of Fabaceae, belonging to the
genera Lotus, Dorycnium, Anthyllis or Coronilla,
see table 1. The stem-mining species of this group
are often sympatric with and feeding within cen-
timeters of leaf-mining species of the subgenus
Trifurcula (Levarchama), which have a very similar
host-range: on Lotus we find T. subnitidella to-
gether with either 7. (L.) eurema (Tutt) or T. (L.)
cryptella (Stainton); on Dorycnium, T. josefklime-
schi occurs together with T. (L.) eurema and on
Antbyllis cytisoides we found T. victoris on the
same plants as T. (L.) anthyllidella Klimesch. Hi-
therto all species of the subnitidella group are found
on a single hostplant genus only.
The egg is deposited on the stem of the host, in
one species on a leaflet. The larvae make relatively
straight gallery mines in the green bark of the stem,
usually changing direction of feeding several times
(figs. 105-107). Larvae feed with their venter to-
wards the epidermis. The full-grown larva quits the
mine through a semicircular slit, and spins a cocoon
on the soil or in leaf-litter.
The life history is still incompletely known. It is
assumed here that most species are univoltine, al-
beit with a long flight period, but bivoltinism is not
excluded, and well possible for T. sxbnitidella and
josefklimeschi. The mediterranean species have
VAN NIEUKERKEN: Trifurcula subnitidella group
Table 1. Hostplants of Trifurcula subnitidella group, all
in family Fabaceae (Papilionaceae).
Hostplant species of Trifurcula
Tribus Loteae
Dorycnium hirsutum (L.) Ser.
D. pentaphyllum Scop.
Lotus corniculatus L.
Anthyllis cytisoides L.
7. T. josefklimeschi
7. T. josefklimeschi
6. T. subnitidella
5. T. victoris
Tribus Coronilleae
Coronilla juncea L. 4. coronillae
been collected as larvae in January, February and
April, and T. subnitidella larvae have been found in
September and October in The Netherlands. Most
specimens of subnitidella and josefklimeschi
emerged within a short period, but T. coronillae and
victoris specimens emerged over a period of several
months.
Although the hostplant and immature stages of
the other five species are unknown, it is tentatively
assumed that they feed on related hostplants (her-
baceous Fabaceae) and make stem-mines as well.
The first assumption is supported by the localities
of several of these species: they have been found in
grassland habitats without any species of broom,
the most likely alternative hosts in this subgenus.
T. iberica and silviae have been found in alpine
meadows, where herbaceous Fabaceae are abundant
and T. puplesisi occurs in western Asia, beyond the
natural range of broom species.
Adults of the subritidella group have been col-
lected by sweeping in grasslands and at light. They
are, however, easily overlooked, and the search for
mines is a more secure way to obtain records.
Distribution
The group has a wide distribution in the western
Palaearctic region, but only T. subnitidella is wides-
pread. The other species center around the mediter-
ranean, currently with the highest number of spe-
cies in Spain (5), southern France (3) and eastern
Austria (3). One species, T. puplesisi, has a much
more eastern distribution: around the Caspian Sea
and in the western part of the Central Asian deserts,
it is the easternmost Palaearctic species of Trifur-
cula s. str. The scarcity of records from Greece and
northern Africa presumably is biassed by collecting
activities.
Checklist of the Trifurcula subnitidella group
Trifurcula Zeller, 1848
subgenus Trifurcula s. str.
211
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
submitidella species group - Underside of forewing with large costal fold,
l. austriaca Sp. n. hiding the distinct androconial patch (figs. 20,
2. luteola sp. n. 58). Hindwing costa with enlarged lobe, a small
3. puplesisi sp. n. group of black scales just behind frenulum (fig.
4. coronillae sp. n. DO io SEEN à 7. T. josefklimeschi
5. victoris sp. n. - Underside of forewing without yellow patch,
6. subnitidella (Duponchel, 1843) but margin of costal fold with row of dark-
griseella Wolff, 1957 brown or black special scales; a row of similar
7. josefklimeschi sp. n. scales along hindwing costa (fig. 60). Pale al-
8. iberica sp. n. PINEISPECICS APE 9. T. silviae
9. silviae sp. n. 2. Yellow patch divided into three well-separated
areas with different tinges: from yolk yellow,
through yellowish white to grey-brown (figs.
Key to the males of the Trifurcula subnitidella 10, 18). Hindwing costa without black scaling
ERO UP Ee OTT si A O EN ei Ge ANNE ei 4. T. coronillae
1. Underside of forewing with a well exposed - Yellow patch with only one colour. Hindwing
patch of yellow(ish) androconial scales near costa with or without black scaling........... 3
wing base (figs. 8-10). Hindwing costa with or 3. Hindwing costa with black scales extending
Wichoutsomeiblacksis cal es rn ee 2 from about % to % (fig. 59)....... 8. T. iberica
Figs. 23-28. Trifurcula austriaca, male genitalia holotype, slide 2591 (except fig. 28). - 23, Capsule, valva outlined; 24,
Juxta; 25, Valva, dorsal aspect; 26, Larger cornuti from paratype, slide Klimesch 757, slightly squashed; 27, Aedeagus,
ventral aspect; 28, Idem, lateral aspect. Scale 0.1 mm.
2112
© coronillae
© iberica
B luteola
A silviae
EA
W austriaca
. Small species (forewing length 2.6-2.9 mm)
Hindwing either with very small group of black
scales near frenulum or without such scales 4
. Hindwing costa with a small row of black scales
just behind frenulum (fig. 57) (difficult to de-
tect). Antenna with 31-36 segments. Gnathos
asymmetrical, with a small keel at right side
(COSO eno 6. T. subnitidella
Hindwing costa completely without black scal-
ing. Antenna with 34-48 segments. Gnathos
either asymmetrical with two keels, or symmet-
GIGAN ate ore Peng aa o Ne ea 5
with 34-37 antennal segments. Androconial
patch (fig. 8) yellowish white, similar scales
also present on hindwing upperside. Aedeagus
with two curved cornuti and one spine-like cor-
OUEUSK(H ONDO) par ne 1. T. austriaca
Larger species (2.8-3.5 mm), with 40-48 anten-
nal segments. Androconial patch a darker yel-
low. Aedeagus without curved cornuti, but with
one or two spine-like and a conical cornutus
. Gnathos asymmetrical, with two large keels
(fig. 51, 52). Aedeagus with two very similar
spine-likercornuti nen 5. T. victoris
Gnathos simple, symmetrical (figs. 30, 37). Ae-
deagus with three differently shaped larger cor-
MUD en Maus art a Ser Een le 7
. Very pale species: scale tips yellow (fig. 2).
Valva almost triangular, gradually narrowing
towards apex (fig. 33). Species from western
Krane tee. tente coco
Darker species: scales with darker tips. Valva
basally with almost parallel margins, distally
suddenly tapering towards apex (fig. 38). Spe-
GIES (HHONIN WIESE JAG i 3. T. puplesisi
VAN NIEUKERKEN: Trifurcula subnitidella group
Fig. 29. Distribution of Trifwrcula spe-
cies, see legend. Mapped on 50 X 50 km
UTM squares.
1. Trifurcula austriaca sp. n.
(figs. 1, 8, 23-28, 29)
Trifurcula n. sp.; van Nieukerken im Kasy 1985: 5.
Type material. - Holotype 8, Austria: Fürbachwiesen,
E. of Gramatneusiedl (Niederöst.), UTM: 33U XP12,
15.v.1968, F. Kasy, Genitalia slide EvN 2591 (NHMW).
- Paratypes, 2 &: same data, but 4.ix.1973, 31.vii.1978
(RMNH, NHMW).
Description
Male (fig. 1). - Forewing length 2.6-2.9 mm,
wingspan 5.6-6.7 mm. Head: frontal tuft yellowish
orange, collar yellowish white. Antenna with 34-37
segments; scape yellowish white, sometimes with a
few dark scales. Forewing and thorax covered with
fuscous tipped scales, mixed with white scales, giv-
ing a grey-brown impression; terminal cilia white
beyond relatively distinct cilia-line. Underside of
forewing with basal patch buff yellow, extending to
‘4 (fig.8). Hindwing with a row of white hair-scales
in stead of costal bristles; basally on upperside with
some yellowish white to buff yellow scales, similar
to forewing underside.
Female unknown.
Male genitalia (figs. 23-28). — Capsule length
330-350 um. Vinculum anteriorly slightly nar-
rowed, truncate; lateral arms near gnathos with
minute spines. Uncus with medial process pointed,
with pair of lateral setae. Gnathos symmetrical,
central element triangular, with wide apical angle;
anterior processes present. Valva length 245-255
um, basally broad, with parallel margins, posterior
half suddenly narrowed towards short pointed tip,
transverse bar of transtilla less than twice the
length of sublateral process. Aedeagus 310-385 um
PAS)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 30-36. Trifurcula luteola , male genitalia. - 30, Capsule, valva outlined, slide 2342; 31, Gnathos, holotype, slide 2503;
32, juxta, holotype, slide 2303; 33, Valva, dorsal aspect, slide 2342; 34, Aedeagus, ventral aspect, holotype, slide 2303;
35, Aedeagus, ventro-lateral aspect, slide 2304; 36, Larger cornuti, holotype, slide 2303. Scale 0.1 mm.
long, with ventral carina posteriorly ending in two
ventrally curved processes; dorsal lobe at right side
indistinct, a large lobe at left side, ventrally enlarged
and strongly sclerotized (fig. 27, 28). Vesica with
three large cornuti (fig. 26): one long pointed (-135
um) and two curved (respectively 85 and 60 um);
further a serrate cornutus lower in vesica, in addi-
tion to the group of needle-like and the small cor-
nuti. Juxta fig. 24.
214
Diagnosis
Externally most similar to sabnitidella, but with-
out black scales near frenulum; austriaca also has
special scales on hindwing upperside. Other species
without black scales are larger and/ or paler
(luteola, puplesisi, victoris). Male genitalia resem-
ble more those of the T. pallidella group than other
species in this group: the two curved cornuti and the
triangular gnathos are diagnostic.
Biology
Immature stages and hostplant unknown. The
adults were all taken at light, from May to Sep-
tember, in a grassland nature reserve, with a very
rich Lepidoptera fauna (Kasy 1985).
Distribution (fig. 29)
Eastern Austria: only from the type locality: the
‘Pischeldorfer Fischawiesen’ (named Fürbach-
wiesen on labels). After finishing this description,
Z. Lastúvka showed me in August 1990 a drawing
of male genitalia undoubtedly belonging to this
species, from a specimen from Czechoslovakia:
Prostéjov Hamry [UTM: 33U XQ48], collected in
1990; this record is added on fig. 29. Possibly a
species with an eastern distribution, like several
other Lepidoptera species occurring in this area.
Etymology
An latinized adjective, from the type locality Aus-
tria.
2. Trifurcula luteola sp. n.
(figs. 2, 29, 30-36)
[Trifurcula pallidella Duponchel
[1963]: 1209. Misidentification.]
partim; Lhomme
Type material. - Holotype &: France, Molières[-sur-
l’Alberte] (Aude), UTM: 31T DH57, 6.viii.1903, Chré-
tien, Genitalia slide EvN 2303 (MNHN). - Paratypes: 4
6. France: 1 ¢ (abdomen missing), data as holotype
(MNHN); 1 6, Arten. [St. Pons, montagne d’Artenac]
(Hérault), UTM: 31T DJ81, 19.vii.1904, Chrétien
(MNHN); 2 3, Nesp. [Nespouls, near St. Pons] (Hé-
rault), UTM: 31T DJ81, 2.viii.1904, (MNHN, RMNH).
Description
Male (fig. 2). - Forewing length (2.4) 3.2-3.4
mm, wingspan (5.4) 7.2-7.6 mm. Head: frontal tuft
yellowish orange, collar paler. Antenna with 40-45
segments; scape white with few yellow scales. Fore-
wing and thorax pale yellowish white, tips of scales
yellow; terminal cilia concolorous. Underside of
forewing with orange yellow basal patch to %, cos-
tal fold with row of brown scales along edge.
Hindwing white, humeral lobe on upperside near
frenulum with a row of yellowish-orange special
scales.
Female unknown.
Male genitalia (figs. 30-36). — Capsule length
370-375 um. Vinculum anteriorly truncate, broad
and short. Uncus with medial process terminally
truncate, not widened, several lateral setae. Gna-
thos symmetrical, central element approximately
triangular, slightly angular or more rounded (figs.
30, 31); long anterior apodemes present, but not
very distinct. Valva length 280-305 um, roughly
triangular, with short pointed tip; transverse bar of
transtilla 2.5-3 X as long as sublateral process. Ae-
VAN NIEUKERKEN: Trifurcula subnitidella group
deagus 405-435 um long, with long, terminally
fringed, ventral carina; vesica with one long,
pointed cornutus (225-250 um), a short conical (75
um), + 10 long needle-like and many small cornuti.
Aedeagal tube with dorsal lobe at right side not very
pronounced, slightly serrate. Juxta fig. 32.
Diagnosis
Externally easy to separate from most other spe-
cies by pale colour and size. Resembles Trifurcula
pallidella (Duponchel), but male easily distin-
guished by yellow patch on forewing underside.
Male genitalia characterized by triangular valva and
triangular, symmetrical gnathos, but see puplesisi.
Biology
Immature stages and hostplant unknown. Adults
collected in July and August.
Distribution (fig. 29)
Only known from Southwest France, in mediter-
ranean region.
Remarks
The four specimens of this species were found
amongst material labelled as Trifurcula pallidella in
the Chrétien collection. Although the species has
only been collected in the beginning of this century,
it is very well possible that it still can be found, but
has until now been overlooked, as was the case with
silviae.
Etymology
A Latin adjective, from /uteolus = yellow, orange,
named after the pale forewing colour.
3. Trifurcula puplesisi sp. n.
(figs. 37-44, 89, 95)
Type material. - Holotype 8: Soviet Union: Turkmen-
istan, Sandykachi (Sandy Katschi), UTM: 41S MA55,
1.v.1986, Puplesis, Genitalia slide EVN 2760 (ZKVV). —
Paratypes: 2 6, 4 Q. Soviet Union: 1 @, 2 Q, same data as
holotype; 2 9, Turkmenia, Central Karakum, env. Ash-
khabad, UTM: 40S FH20, 4.vi.1988, R. Puplesis (ZK VV,
RMNH); 1 4, [Derbent (Dagestan), UTM: 39T TG76],
10.vii.[1872], [ Nacht bei Ligt’], Christoph, ‘473’, Zeller
Coll., Walsingham collection (BMNH).
Excluded from type-series (probably conspecific): 1 8,
Krasnoarmeysk (Sarepta), UTM: 38U MX77, Christoph,
‘397’ [red label] (BMNH).
Description
Male. - Forewing length 2.9-3.7 mm, wingspan
6.2-8.0 mm. Head: frontal tuft pale wish orange to
almost white, collar white. Antenna with 46-48
segments; scape white. Forewing and thorax
greyish brown, scales with dark tips, except along
dorsum; terminal cilia white. Underside of fore-
wing with a small pale yellow basal patch to \/s.
215
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Hindwing white, humeral lobe on upperside near
frenulum with some yellow special scales, similar to
those on forewing underside; costa basally with row
of white hair-scales.
Female. - Forewing length 3.2-3.3 mm (3.25 +
0.04, 4), wingspan ca 7.3 mm. Antenna with 43-45
segments.
Male genitalia (figs. 37-43). — Capsule length
350-410 um. Vinculum wide and short, anteriorly
broadly rounded. Uncus with medial process termi-
nally truncate or bifurcate (figs. 37, 39). Gnathos
symmetrical, central element broadly rounded, an-
terior apodemes not evident. Valva length 275-295
um, broad, margins in basal half approximately
parallel sided, tip slightly curved, pointed; trans-
verse bar of transtilla approximately twice as long
as sublateral process. Aedeagus 350-435 um long,
with long, terminally fringed, ventral carina and a
dorsal lobe at right side, with serrate margin. Vesica
with one long, curved, pointed cornutus (155-190
pm), a short conical (65-100 um), a long irregular
cornutus, seeming composed of several cornuti and
almost hidden amongst the long needle-like cor-
nuti; many small cornuti present. Juxta fig. 40.
Figs. 37-43. Trifurcula puplesisi, male genitalia. - 37, Capsule, valva outlined, slide 2761; 38, Valva, dorsal aspect, slide
2761; 39, Slightly different uncus of holotype, slide 2760; 40, Juxta, slide 2761; 41, Cornuti, dorsal aspect, slightly squashed,
slide BM 25634; 42, Aedeagus, ventral aspect, part of cornuti, protruding from aedeagal tube also shown, slide 2761;
43, Aedeagus, lateral aspect, holotype, slide 2760. Scale 0.1 mm.
216
Female genitalia (figs. 89, 95). - Terminal seg-
ments relatively broad; T8 with about 9-11 setae
and several spines, broad anal papillae with 28-37
setae. Signa long, 525-630 um. Ductus spermathe-
cae with 34 coils.
Diagnosis
Males resemble other species without black scal-
ing and simple yellow patch, in particular austriaca,
which is smaller and has fewer antennal segments
and /wteola, which is usually much paler and has a
more triangular valva. Female recognized by large
number of setae on anal papillae and relatively long
signa.
Biology
Hostplant and immature stages unknown. The
adults were taken in steppe area, almost desert, in
May, June and July.
Distribution (fig. 44)
Caspian Sea area, just inside Europe and southern
Turkmenia.
Remarks
The specimen from the Christoph collection, la-
belled ‘Sarepta’ is tentatively assigned to this spe-
cies, but excluded from the type series, since it is
much paler and shows slight differences in genita-
lia, such as the form of the juxta.
The paratype from the Zeller/Walsingham col-
lection, also collected by Christoph, has only been
labelled with: ‘10/7’ [black ink on green, handwrit-
ing] / ‘473 [round label, black ink on green, handw-
riting] / ‘Christoph’ [black ink on white, handwrit-
ing], and the usual Walsingham collection labels
with no. 101450. The locality has been traced from
correspondence from Christoph to Zeller, now
present in the Entomology Library, BMNH. Only
in one letter with lists of specimens, sent by Chris-
VAN NIEUKERKEN: Trifurcula subnitidella group
Fig. 44. Distribution of Trifurcula puplesisi,
mapped on 50 X 50 km UTM squares.
toph to Zeller for identification, a number as high
as ‘473’ could be found. It is assumed that this refers
to the specimen in question. The letter is dated
‘Sarepta Den 12 Marz 1873’ and bears the number
382. Under the number ‘473’ in the list ‘Determi-
nanda’ is written ‘Derbent, Nacht b Ligt’. It is also
assumed that the specimen was collected in July
1872, since the letter apparently refers to the most
recent sending, collected in the previous year. Note
further that the locality Derbent, a town on the
Caspian Sea, is sometimes confused in lepidopteran
literature with the locality ‘Derbend’ in Iran, North
of Teheran, where recently various lepidopterists
collected. Christoph often travelled via Derbent,
when he crossed the Caspian Sea towards Turkmen-
istan or northern Iran (Christoph 1877), and as far
as I know never collected in the Iranese ‘Derbend’.
Etymology
A noun in genitive, named in honour of its col-
lector, Dr. Rimantas Puplesis, a Lithuanian special-
ist of central and east Asian Nepticulidae, who
kindly trusted me his material for description.
4. Trifurcula coronillae sp. n.
(figs. 3, 10, 18, 19, 29, 45-50, 90, 96, 101, 106)
Type material. - Holotype @: Spain, Sierra Blanca, 6 km
N Marbella, El Mirador (Malaga), 800 m, UTM: 30S
UF34, 5-9.11.1984, Coronilla juncea, from stem-mines, el.
13.vii.1984, EvN no. 84046, E. J. van Nieukerken, Geni-
talia slide EVN 2592 (RMNH). - Paratypes: 15 6, 11 9,
30 larvae. Spain: 4 8, 4 9, 7 larvae, data as holotype, e.l.
31.v.1984- 25.viii.1984, (RMNH); 13 larvae, 5 km S Istán,
road to Embalse de la Concepción (Málaga), 150 m, UTM:
30S UF2545, 17.1.1988, Coronilla juncea, stem-mines,
EvN no. 88057, van Nieukerken & Richter (RMNH); 4
8, El Saler (Valencia), UTM: 30S YJ26, 22.iv.1981, C.
Gielis (RMNH, coll. Gielis); 2 larvae, Rodalquilar, 5 km
SW Las Negras (Sierra del Cabo de Gata) (Almería), 120
m, UTM: 30S WF8579, 8.1.1988, Coronilla juncea, stem-
2417,
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 45-50. Trifurcula coronillae, male genitalia holotype, slide 2592 (except 48, 49). - 45, Capsule, valva outlined; 46,
Valva, dorsal aspect; 47, Juxta; 48, Large cornuti, separately drawn, slide 2593; 49, Aedeagus, ventral aspect, slide 1883;
50, Aedeagus, lateral aspect. Scale 0.1 mm.
mines, EvN no. 88005, van Nieukerken & Richter
(RMNH); 2 6, 4 9, Sierra Blanca, El Garopala, 3 km
NNW Ojén (Malaga), 670 m, UTM: 30S UF3250,
15.1.1988, Coronilla juncea, stem-mines, e.l. 6.vi.1988-
22.v111.1988, EvN no. 88051, van Nieukerken & Richter
(RMNH); 4 8, 3 9, 8 larvae, Sierra Blanca, Puerto de
Marbella (El Mirador), 3 km W Ojén (Málaga), 900 m,
UTM: 30S UF3048, Coronilla juncea, 15.1.1988, el.
30.v.1988-30.vii.1988, EvN no. 88047, van Nieukerken &
Richter (RMNH); 1 8, road to Istan (Málaga), 400 m,
UTM: 30S UF24, 25.vi.1975, E. Traugott-Olsen (ZMUC).
Excluded from type series: 2 & (abdomina lost), data
as holotype, el. 1.viii.1984 (RMNH).
Description
Male (fig. 3). - Forewing length 2.6-3.2 mm (2.88
+ 0.17, 18), wingspan 5.6-6.7 mm. Head: frontal
tuft yellowish orange to orange , collar slightly paler
to almost white. Antenna with 41-48 segments
(44.0 + 2.0, 12); scape white, often with scattered
218
brown scales. Forewing and thorax covered with
brown tipped scales, distal 4 dark, edge darker,
giving the wing a greyish brown impression; white
scales sometimes exposed at dorsum, terminal cilia
yellowish white, cilia line indistinct. Underside of
forewing with large patch, extending from base to
%, more or less distinctly divided into three areas
with different colour: basally deep yellow, middle
part buff yellow, terminally olive brown (figs. 10,
18, 19). Hindwing greyish, basal area with white
lamellar scales.
Female. - Forewing length 2.4-3.2 mm (2.91 +
0.23, 11), wingspan 5.4-6.4 mm. Antenna with 35-
42 segments (39.8 + 2.2, 10).
Male genitalia (figs. 45-50) - Capsule length 340-
355 um. Vinculum anteriorly truncate. Uncus with
medial process truncate. Gnathos asymmetrical,
with a longitudinal keel, slightly left of middle on
ventral surface, posterior margin rounded. Valva
length 250-270 um, with almost parallel margins,
suddenly narrowed in terminal quarter towards
pointed tip, transverse bar of transtilla twice as long
as sublateral processes or longer. Aedeagus 360-390
um long, with ventral carina bifurcate; aedeagal
tube with spatulate tip ventrally, dorsal lobe not
pronounced at right side, but with strong medial
spine-like process, curved towards right side, which
in normal preparations is almost inseparable from
the large cornuti: there appear to be three in stead
of two. Vesica with two large spine-like cornuti
(one somewhat irregular) and a conical cornutus;
long needle-like and many smaller cornuti present.
Juxta fig. 47.
Female genitalia (figs. 90, 96). Terminal seg-
ments broad and rounded; T8 with 6-8 setae and
several scales; anal papillae with 24-30 setae. Duc-
tus spermathecae with 3% to 3/2 coils. Signa 420-
470 um (ventral) and 460-520 um (dorsal) long.
Final instar larva (fig. 101). Yellow, elongate.
Headcapsule 325-355 um long (341 + 10.7, 5),
1.06-1.13 (1.09 + 0.02, 5) X as long as wide. Spin-
osity: prothorax ventrally with narrow band poste-
riorly, meso- and metathorax ventrally with a band
along anterior margin and very few spines on calli.
Abdominal segments 1-9 with sparse spinosity ven-
trally and laterally up to the spiracles, spines near
anterior margins smaller; segment 10 with small
group of spines anterior of setae.
Diagnosis
Male easily recognized by the androconial patch
with three different colours and absence of black
scaling on hindwing. Male genitalia unique by
shape of gnathos and cornutus-like process on ae-
deagus. Female very similar to josefklimeschi, but
separated by the blunt abdominal point, versus the
slightly pointed condition in gosefklimeschi. Fe-
males of victoris are paler, and have less antennal
segments, females of subritidella are darker and
have also less antennal segments.
Biology
Hostplant. - Coronilla juncea L., a shrub up to 1
m, with rush-like stems, slightly resembling a
broom.
Life history. - The egg is deposited on an inter-
nodium of the smooth rush-like stem of the host-
plant, and is conspicuous. The larva makes a very
conspicuous gallery mine in the green bark, in
which the living larva can easily been seen (fig.
106). The mine often starts contorting, frequently
encircling the stem; the larva often feeds downward
first, but later changes feeding direction one or
more times, the mine usually ends in upwards di-
rection. The mine is filled with greenish brown
frass. There are often numerous mines in a single
stem. The brown cocoon is made in soil or on leaf
VAN NIEUKERKEN: Trifurcula subnitidella group
litter. Larvae have been collected in January and
February, but many mines were already vacated in
that period, so that larvae probably start mining in
autumn. Adults emerged over a very long period,
from May to August. The only adults taken in the
field were collected in April and June. The species
is most likely univoltine.
Distribution (fig. 29)
Along mediterranean coast of Spain, to be ex-
pected elsewhere with its host, which occurs from
West Yugoslavia to Portugal (Heywood & Ball
1968).
Etymology
A noun in genitive case, from Coronilla, the ge-
neric name of the hostplant.
5. Trifurcula (Trifurcula) victoris sp. n.
(figs. 4, 51-56, 76, 91, 97, 102)
Type material. - Holotype @: Spain: El Pozo de los
Frailes, 3 km N San José (Almeria), UTM: 30S WF7972,
8.1.1988, Anthyllis cytisoides, stem-mines, e.l. 4.iii.1989,
EvN no. 88003, van Nieukerken & Richter (RMNH). —
Paratypes: 2 6,6 9, 4 larvae. Spain: 2 8,6 9, 3 larvae, same
data as holotype, el. 29.ii.1989-30.v.1989 (RMNH); 1
larva, 3 km NE Nijar, Cerro de Movillas (Almeria), 400
m, UTM: 30S WF7393, 9.1.1988, Anthyllis cytisoides,
stem-mines, EvN no. 88007, van Nieukerken & Richter
(RMNH).
Description
Male (fig. 4). - Forewing length 3.0-3.5 mm,
wingspan 6.5-7.2 mm. Head: frontal tuft yellowish-
orange to ferruginous, collar paler, yellowish white.
Antenna with 40-42 segments; scape white, with
some darker scales, flagellum brown. Forewing and
thorax relatively pale, appearing pale brown-grey:
scales with distalmost tip (margin only) brown, few
white scales at tornus; terminal cilia white beyond
more or less distinct cilia-line. Underside of fore-
wing with deep yellow patch of about ‘4 forewing
length. Hindwing grey, humeral lobe well deve-
loped, on upperside with yellow scales along mar-
gin, black scaling absent.
Female. - Forewing length 2.9-3.3 mm (3.04 +
0.12, 6), wingspan 6.5-7 mm. Antenna with 32-35
segments (33.5 + 1.3, 4). Ovipositor with short
point.
Male genitalia (figs. 51-56) — Capsule length 425-
440 um. Vinculum anteriorly truncate or rounded.
Uncus with medial process strongly widened, trun-
cate. Gnathos very asymmetrical, variable, always
with two longitudinal keels of different length,
leaving ventrally excavation in which aedeagal tip
fits (dashed in fig. 51), anterior apodemes present.
Valva length 285-290 um, approximately triangu-
lar, dorsally with a row of 4-5 spines along conti-
nuation of lateral arm of transtilla, transverse bar
219
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 51-56. Trifurcula victoris, male genitalia. - 51, Capsule, valva and tip of aedeagus outlined, slide 3064; 52, Gnathos
of holotype, showing variation, slide 2743; 53, Valva, dorsal aspect, slide 3064; 54, Large cornuti, separately drawn, slide
3064; 55, Aedeagus and juxta, ventral aspect, slide 3064; 56, Aedeagus and juxta, lateral aspect, holotype, slide 2743. Scale
0.1 mm.
of transtilla about twice as long as sublateral pro-
cesses. Aedeagus 395-435 um long, with ventral
carina spatulate, dorsal lobe at right side conspicu-
ous, margin serrate; vesica with two large and very
similar cornuti, one joined to smaller cone-shaped
cornutus; group of spine-like cornuti very much
reduced in size and number, only a few present at
basis of both large cornuti; numerous small cornuti
present. Juxta fig. 55.
Female genitalia (figs. 91, 97). - Terminal seg-
ments forming slightly pointed abdominal tip; T8
220
with 8-10 setae and some scales; anal papillae with
21-29 setae. Ductus spermathecae with 4 coils.
Signa 465-530 um long.
Final instar larva (fig. 102). - Yellow, elongate.
Head capsule 385-435 um, 1.03-1.05 X as long as
wide. Spinosity: prothorax ventrally with narrow
band of minute spines posteriorly, meso- and me-
tathorax ventrally without spines. Abdominal seg-
ments 2-9 with microspines ventrally, hardly vis-
ible, segment 10 with small group of spines
anterior of setae.
Diagnosis
Male recognized by uniformly coloured yellow
patch, absence of black scaling and relatively pale
forewings. T. /uteola is paler, with yellow scale tips
instead of brown or grey. Male genitalia character-
ized by asymmetrical gnathos with two keels and
reduction of spine-like cornuti. Female may be con-
fused with coronillae, which is darker; victoris also
has more pointed anal papillae.
Biology
Hostplant. - Anthyllis cytisoides L., a broom-like
leafy shrub of 0.5-1.5 m, with velvety hairy stems
and leaves.
Life-history. - Egg deposited on stem. The larva
makes a long gallery in the bark, which is difficult
to see, because of the dense indumentum of the
stem. It could best be seen because of the swelling
and the slight discoloration, especially in the case of
old mines. Larva not visible in its mine. The mines
were actually collected on stems, heavily infested by
gall-forming coccids. Larvae collected in January,
adults emerged from February to May. One female
emerged from leaf-litter, which had been collected
from underneath the hostplants.
Distribution (fig. 76)
Only known from the extremely dry region in the
southeast part of the province of Almeria, Spain.
Despite search in other sites along the Spanish
southcoast, where the host is common, no more
mines could be found.
Etymology
A noun in genitive case, from latin victor, named
after my son Victor Alexander.
6. Trifurcula (Trifurcula) subnitidella
(Duponchel)
(figs. 5, 9, 11-17, 57, 61-68, 92, 98, 103, 107)
Elachista subnitidella Duponchel, [1843]: 326, pl. 77: 8.
Lectotype @ [designated by van Nieukerken & Johans-
son 1987: 471]: [Austria, Vienna region], Duponchel
coll., Genitalia slide EvN 2522 (MNHN) [examined].
Trifurcula griseella Wolff, 1957: 21. Holotype 4: Den-
mark, Asserbo, 30.v.1954, N. Wolff, genitalia slide
NW 1797 (ZMUC) [Synonymized by van Nieukerken
& Johansson 1987: 471] [examined].
Nepticula subnitidella (Duponchel) Zeller 1848: 305
[redescription, mistaken identity], Joannis 1915: 127
[identity], Rebel 1901: 221 [catalogue], Meess 1910:
474 [listed].
Trifurcula subnitidella (Duponchel) van Nieukerken &
Johansson 1987: 471 [selection lectotype], 1990: 273
[description, NW Europe].
Trifurcula griseella Wolff, Bradley 1962: 174, fig. 4
Ireland, male genitalia], Bjorn & Pallesen 1971: 111
[Denmark], Bradley et al. 1972: 3 [Britain], Svensson
1974: 171 [Sweden], Emmet 1975: 39-42 [England,
Scotland], Karsholt & Nielsen 1976: 18 [Denmark],
Emmet 1976: 209 | description, British Isles], Svensson
VAN NIEUKERKEN: Trifurcula subnitidella group
1980: 85 [Sweden], Buhl et al. 1983: 120 [Denmark],
Svensson 1983: 65 [Sweden], Kasy 1983: 5 [ Austria],
Buhl et al. 1984: 3 [Denmark], Svensson 1985: 81
[Sweden], Kasy 1985: 5 [ Austria], Karsholt 1985: 45
[catalogue Denmark], van Nieukerken 1986a: 16
[checklist], Bradley & Fletcher 1986: 2 [list], Prose
1987: 49 [West Germany: Bavaria], Svensson 1987: 3-
3 [catalogue Sweden], Buhlet al. 1987: 100 [Denmark],
Buhl et al., 1988: 101 [Denmark].
Description
Male (fig. 5). - Forewing length 2.0-2.6 mm (2.37
+ 0.17, 44), wingspan 4.2-5.8 mm. Head: frontal
tuft yellow-ochre to fuscous, variable; collar similar.
Antenna with 31-36 segments (33.4 + 1.3, 40);
scape white, sometimes with few brown scales.
Forewing dark greyish-brown to fuscous, slightly
irrorated by dark tipped scales, no white spots pres-
ent; underside with distinct basal patch of deep
yellow scales-near costa reaching to % (figs. 9, 11-
13). Hindwing: grey, underside along costa directly
behind frenulum with a distinct short row of black
lamellar scales (figs. 14, 15, 57). Abdomen dark
grey, with yellowish grey tufts.
Female. - Forewing length 2.0-2.4 mm (2.22 +
0.16, 9), wingspan 4.4-5.4 mm. Antenna with 27-32
segments (30.4 + 1.7, 9).
Male genitalia (figs. 61-67). — Capsule length
330-390 um (349.6 + 14.8, 15). Vinculum ante-
riorly rounded. Uncus slightly widened, truncate at
tip. Gnathos asymmetrical, central element with an
anteriorly curved keel, ending in a pointed process
at right side; lateral arms with long and narrow
anterior apodemes. Valva length 250-289 um
(264.2 + 9.0, 15), basally with almost parallel mar-
gins, narrowed in middle towards pointed tip. Ae-
deagus 330-365 um (344.0 + 14.0, 15), with ventral
carina bifid; aedeagal tube with spatulate tip, dorsal
lobe in middle, or slightly on right side, with serrate
margin. Vesica with one very long straight, or
slightly curved cornutus, with a conical cornutus
joined to its basis, and a pointed cornutus with
serrations; less then 10 spine-like cornuti and many
small cornuti present. Juxta fig. 62.
Female genitalia (figs. 92, 98). - Terminal seg-
ments rather broad; T8 with 3-6 setae and some
scales; anal papillae with 11-20 setae each. Ductus
spermathecae with 314 convolutions. Signa 300-355
um (dorsal) and 350-365 um (ventral) long.
Final instar larva (fig. 103). - Long and slender,
deep yellow. Head capsule 325-345 um, 1.13-1.16 X
as long as wide, distinctly narrower than in other
three described species. Spinosity: prothorax ven-
trally with narrow band posteriorly, meso- and me-
tathorax with spines between D1 seta and calli.
Abdominal segments 1-9 with sparse spinosity ven-
trally and laterally up to the spiracles, spines near
anterior margins smaller; segment 10 with small
group of spines anterior of setae.
(NO)
159]
A
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 57-60. Trifurcula males, forewing and hindwing undersides, showing position of black scales, yellow patch and
costal fold. - 57, T. subnitidella, Netherlands, Katwijk; 58, T. josefklimeschi, paratype, Austria, Hundsheimer Berg; 59,
T. iberica, hindwing only, paratype, Spain, Prullans; 60, T. silviae, paratype, France, Ceillac. Scale 0.3 mm.
Diagnosis
Male subnitidella differs from other species in
the group by the combination of a simple yellow
patch and a group of black scales just behind fren-
ulum; further it is one of the smaller and darker
species, with less antennal segments than most
other species (36 maximum). The male genitalia
are well characterized by the gnathos process. Fe-
males are smaller and darker than the other des-
cribed females, and have less antennal segments.
Biology
Host plant. - Lotus corniculatus L., a common
perennial herb of grasslands all over Europe, pos-
sibly also on other Lotzs species.
Life history. - The egg is conspicuous and dep-
osited on the stem of the host, usually at the base
222
of a leaf-stalk (fig. 107), frequently near the stem
base. The mine is a long gallery in the bark of the
stem (fig. 107): the larva is first mining down for
0.5-2 cm, then going up the stem ina rather straight
line, or partly encircling the stem, occasionally go-
ing down again in the last part of the mine. The
mine may reach a total length of 6 to 9 cm. The
mine is at first narrow, reddish brown, with straight
edges, but later becomes as wide as the stem, with
irregular margins, becoming silvery white in fresh
mines. Frass deposited in midline, brown, not al-
ways well visible. Larva well visible in mine, ap-
pearing as a slight swelling. The stems of the host
are not killed by the mining activity. The larva quits
the mine through a semicircular slit, and spins a
cocoon in the soil or on leaf-liter.
VAN NIEUKERKEN: Trifurcula subnitidella group
62
65
Figs. 61-67, Trifurcula subnitidella, male genitalia, slide 2501 (slide 2745 for fig. 62). - 61, Capsule, valva outlined; 62,
Juxta; 63, Valva, dorsal aspect; 64, Aedeagus, ventral aspect; 65, Idem, dorsal lobe, dorsal aspect; 66, Large cornuti,
separately drawn; 67, Aedeagus, lateral aspect. Scale 0.1 mm.
Larvae have been taken in September and Oc-
tober, adults fly from May (March in Tunisia) to
early September. Reared adults emerged in May.
Voltinism not yet clear from these data, clear peaks
in flight are not apparent. More data from single
localities are needed, and bivoltinism can only be
proved by finding larvae and rearing in early
summer.
In northern Europe T. submitidella is almost ex-
clusively found on limestone grasslands (down-
land) and coastal dunes, in southern Europe it is
found in many habitat types. The species might be
vulnerable to habitat loss in more northern parts of
its occurrence.
Distribution (fig. 68)
Widespread in Europe, northward to 60° N,
southward to the northern border of the Sahara in
Tunisia and eastward to the Crimea and Asia minor.
Not yet recorded from Norway, Belgium, Luxem-
bourg, Switzerland, Poland, Hungary, Albania and
Portugal, nor from any of the large mediterranean
islands. The species is here for the first time in
detail recorded from Finland, The Netherlands,
Germany, Czechoslovakia, France, Spain, Italy, Ru-
mania, Bulgaria, Yugoslavia, Greece, Turkey, Soviet
Union (Estonia and Ukraine) and Tunisia.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Remarks
T. subnitidella is slightly variable in colour, spec-
imens from northern Europe tend to be darker than
those from southern Europe.
The figure of the male genitalia in Wolff's des-
cription of T. griseella (1957), has been published
as mirror image.
In North America, a striking case of convergence
has been described by Wagner (1987): his species
Microcalyptris lotella was reared from very similar
stem-mines in Lotus, and, moreover as adult male
it has a yellowish patch on the forewing underside,
very similar to that of sabnitidella. Also lotella has
been found in a habitat that in Europe could har-
bour subritidella: coastal dunes. Yet Microcalyptris
Braun (synonymized with Acalyptris Meyrick by
van Nieukerken 1986a) is a completely different
genus, only distantly related to Trifurcula.
Material examined. - 115 8, 14 9, 5 larvae (including
type-material cited above). - Austria: 1 8 [genitalia slide
MV13111 only], Apetlon, Seewinkel (Burgenland), UTM:
33T XN38, 9.viit.1963, E Kasy (NHMW); 1 5, Fürbach-
wiesen, East of Gramatneusiedl (Niederöst.), UTM: 33U
XP11, 23.vii.1981, E Kasy (NHMW); 1 6, Hundsheimer
Berg S. ( (Niederöst.), UTM: 33U XP43, 8.vil. 1980, F. Kasy
(NHMW); 1 4 [photo genitalia slide], Linz, surroundings
(Oberöst.), UTM: 33U VP44, 18.vii.1934, J. Klimesch
(coll. Klimesch). — Bulgaria: 1 8, Nesebur (Nessebar),
UTM: 35T NHS2, 23.viii-4.ix.1962, J. Soffner (DEIC). —
Czechoslovakia: 2 &, Detkovice (Moravia), UTM: 33U
XQ57, 2.vi and 4.vii.1989, A. LaStuvka (coll. LaStuvka). —
Denmark: 1 9, Asserbo, UTM: 33V UC11, 10.vi.1964, N.
L. Wolff (ZMUC); 6 8, Laesö, Bovet ( (NE), UTM: 33V
PJ25, 8,9 and 13.vii.1982, 30.viand 8.vii.1983, O. Karsholt
(ZMUC); 9 6, 5 9, Laesö, Höjsandet (NE]), NINE 33V
PJ25, 1 and 6.vii.1983 O. Karsholt (ZMUC); 4 8, Laesö,
Nordmarken (NEJ), UTM: 33V PJ25, 5, 6 and 9.vii.1983,
O. Karsholt (ZMUC). - Finland: 2 4, Virolahti (EK),
UTM: 35V NH30, 27.vi.1989 [netted at seashore over
Lotus corniculatus |, J. R. Kaitila (RMNH). - France: 1 4,
Cannes (Alp. Mar.), UTM: 32T LP42, iv.1881, Wal-
singham (BMNH); 1 8, Col de Soubeyrand (Drôme), 994
m, UTM: 31T FK81, 2.viii.1986, H. W. van der Wolf (coll.
van der Wolf); 1 @, La Veuve, Digne, UTM: 32T KP78,
12.viii.1901, Chrétien (MNHN); 1 g, Nesp. [probably
Nespouls, near St. Pons] (Hérault), UTM: 31T DJ81,
2.viii.1904, (MNHN). - Germany (West): 2 4, Bamberg:
Tütschengereuth (Bayern), UTM: 32U PA22, 4.viii.1978,
G. Derra (coll. Derra). - Germany (East): 2 6, Blanken-
burg, Muschelkalk UTM: 32U PBS1, 29.v.1965,
10.viii. 1989, H. Steuer (coll. Steuer). - Great Britain: 1 Ó,
Mickleham (Surrey), UTM: 30U XB88, 23.vi. 1856, grass,
H. T. Stainton (BMNH); 1 6, Mickleham, Headley Lane
(Surrey), UTM: 30U XB88, 10.vi.1857, (BMNH). -
Greece: 1 g, Kavisos (Evro), 100 m, UTM: 35T ME, 22-
23.viii.1985, A. Moberg (NHRS). - Italy: 2 6, Almese,
surroundings (Torino), 350 m, UTM: 32T LQ79, 16 and
20.v.1979, U. Parenti (RMNH, coll. Parenti); 1 8, Asti,
Boschi di Valmanera, 124 m, UTM: 32T MQ37, 1.v.1970,
G. Baldizzone (GBA); 2 8, Asti, Valmanera (Asti), UTM:
32T MQ37, 28.iv.1968, 26.viii.1968, G. Baldizzone
(RMNH, coll. Passerin- d’Entreves); 1 6, Baia Domizia
(Caserte), UTM: 33T UF55+, 25.vii.1972, R. Johansson
(coll. Johansson): 1 8, Cardona (Alessandria), 300 m,
224
UTM: 32T MQ39, 19.v.1975, G. Baldizzone (GBA); 2 è,
Lucotena (Firenze), 500 m, UTM: 32T PP93, 1 l.viii.1982,
J. Kuchlein (RMNH, coll. Kuchlein); 1 @, Monti Aurunci,
S km N Ieri (Latina), 600 m, UTM: 33T UF77, 24-
30.v. 1969, R. Den (coll. Johansson); 1 8, Poggio di
Casasco (Alessandria), 300 m, UTM: 32T NQ06,
3.vi. 1978, G. Baldizzone (GBA), 2 a, Val Susa (Piemonte),
Villardora (Torino), 500 m, UTM: 32T LQ79, 26.v.1983,
G. Bassi (coll. Bassi). - Netherlands: 1 &, Katwijk, 2 km
N: dunes (Zuid-Holland), UTM: 31U ET9686, 16.x.1988,
Lotus corniculatus, stem-mines, e.l. 21.v.1989, EvN no.
88175, van Nieukerken & Richter (RMNH); 1 @, Kun-
rade, Kunderberg (Limburg), UTM: 31U GS0739,
02.vii.1983, G. R. Langohr (coll. Langohr); 2 @, 2 9, 3
larvae, Kunrade: Kunderberg W. (Limburg), UTM: 31U
GS0739, 5.x.1988, Lotus corniculatus, stem-mines, el.
10.v.1989-18.v.1989, EvN no. 88156, van Nieukerken &
Richter (RMNH); 1 6, St. Pietersberg, Cannerbos (Lim-
burg), UTM: 31U FS8733, 18-19.vii.1950, [22.3-0.3 hrs]
(RMNH); 1 6, St. Pietersberg, Zonneberg (Limburg),
UTM: 31U FS8934, 20.vii.1950, [21-22 hrs] (RMNH); 1
d, 19,2 larvae, Wijlre, 1 km SE: railway banks (Limburg),
UTM: 31U GS0534, 5.x.1988, Lotus corniculatus, stem-
mines, e.l. 11.v.1989-15.v.1989, EvN no. 88162, van Nieu-
kerken & Richter (RMNH); 1 4, 1 9, Vrakelberg, 2km
E. of Wijlre, UTM: 31U GS0537, 22.1x.1989, Lotus
corniculatus, stem-mines, el. 1-5.v.1990, E. J. van Nieu-
kerken (RMNH); mines, Noord-Bakkum, 3 km N Castri-
cum (Noord-Holland), dune meadows, UTM 31U
FU1226, 6.x.1989, Lotus corniculatus, E. J. van Nieu-
kerken. - Rumania: 1 g, Gusterita near Sibiu (Hammers-
dorf near Hermannstadt), UTM: 35T KL87, 12.v.1920,
(NHMW); 1 g, Sibiu (Hermannstadt, Pralb.), UTM: 35T
KL87, 18.v.1922 (NHMW). - Spain: 1 6, Beuda (Pyr.
orient.), UTM: 31T DG77, 14.vii.1967, E. Arenberger
(LNKD); 1 4, Biescas (Huesca), UTM: 30T YN12,
1.viii.1989, C. Gielis (coll. Gielis); 3 &, Puerto de Mora
(Granada), 1350 m, UTM: 30S VG52, 22.vii.1986, C. Gie-
lis (RMNH, coll. Gielis); 1 &, San Miguel de Valero N, 3
km S Linares de Riofrio (Salamanca), 850 m, UTM: 30T
TK59, 2.vili.1986, at light ML, Quercus pyrenaica forest
& heathland, EvN no. 86091, EJ. v.Nieukerken &
S.Richter (RMNH); 1 6, Sierra de Marbella, El Mirandor
(Malaga), 700 m, UTM: 30S UF34, 21.vii.1981, E. Trau-
gott-Olsen (ETO); 1 8, road Baza-Benamaurel, 15 km
from Baza (Granada), UTM: 30S WG25, 16.vii.1987, Bal-
dizzone & Traugott-Olsen (GBA); 1 8, road to (Camino
de) Ojen (Málaga), 150 m, UTM: 30S UF34, 25.vi.1983,
E. Traugott-Olsen (ETO). - Sweden: 2 &, 1 > Byrum
Sandvik (Öland), UTM: 33V XD1742, 20.vi.1978, B.
Bengtsson (coll. Johansson, ZMUC); 1 © [slide only],
Kinnekulle (Vg), UTM: 33V VEO9, 29-30.v.1968, I.
Svensson (coll. Svensson); 1 @, Klagshamn (Sk), UTM:
33V UB65, 7.viii.1982, R. Johansson (coll.Johansson). —
Soviet Union: 1 8, Dobroe, Krasnolesje [near Simfe-
ropol] (Krim, Ukraine), UTM: 36T XQ06, 10.v.1982,
Zaguljaev (ZMAS); 1 g, Rakvere (Raustfer) [near Tal-
linn] (Estonia), UTM: 35V MF68, 8.vi.1890 (ZMAS); 1
3, Sevastopol, Inkerman (Krim, Ukraine), UTM: 36T
WQ44, V. Pliginski (ZMAS). - Tunisia: 1 g, Nefta, UTM:
32S LC94, 14-16.111.1986, Zool. Mus. Copenh. (ZMUC). -
Turkey: 2 &, Ankara, 10 km NW Kizilcahaman, 1150-
1250 m, UTM: 36T VK68, 6-7.viii.1989, Fibiger & Esser
(ZMUC). - Yugoslavia: 1 6, Herceg Novi, Igalo, UTM:
34T BN90, 1.v.1938, H. G. Amsel (LNKD); 16 6, Krk,
Draga Baska (Kroatia), UTM: 33T VK78, 25 and
27.vii.1976,3.viii.1976, l.viii.1977, 3.vitt.1978, 5.v111.1985,
30.vii.1986, 15.viii. 1988, G. Baldizzone (GBA, RMNH); 6
VAN NIEUKERKEN: Trifurcula subnitidella group
Fig. 68. Distribution of Trifurcula subnitidella, mapped on 50 X 50 km UTM squares.
8, 2 9, Krk, Misucaynica (Kroatia), UTM: 33T VK78,
18.viii. 1973, 30.vii. 1976, 6.viii.1976, 4 and 9.viii.1986, G.
Baldizzone (GBA, RMNH); 1 4, Krk, Paprata/Vinder
(Kroatia), UTM: 33T VK79, 16.viii.1978, G. Baldizzone
(GBA); 1 8, Krk, Soline (Kroatia), UTM: 33T VK69,
11.viii.1976, G. Baldizzone (GBA); 8 4, Krk, road Krk-
Vrbnik (Kroatia), UTM: 33T VK78, 2.viii.1987,
23.v11.1988, 3.vitt.1988, G. Baldizzone (GBA, RMNH)
Additional records. — Czechoslovakia: Celechovice
(UTM: 33U XQ78), Prostéjov Hamry (33U XQ48), Ko-
zarovce (34U CU15) (Z. Laötúvka in litt.). - Denmark:
Melby Overdrev (Buhl et al. 1983), Korevle (Buhl et al.
1984), Glatved (32V PH14)(Buhlet al. 1987a), Oster Vra
(32V NJ75)(Buhl et al. 1988). - Great Britain: vice coun-
a 1, ORTO MEI AS IGLO MG ORS D Pan dell03
(Emmet in litt. 1987). - Sweden: Hall. Vallda Sandö
(PJ7674), Sm. Ryningsnäs (WD5647), Ol. Algutsrum
(WC9284), Gel. Tingstäde (CK5802), Sdm. Osmo
(XF6739), Upl. Fysingen (XG6407) (Svensson in litt.
1987).
7. Trifurcula josefklimeschi sp. n.
(figs. 6, 20-22, 58, 69-76, 93, 99, 104, 105)
[ Trifurcula orientella Klimesch, 1953: 168, 169 partim: all
specimens from Austria. Misidentification. ]
Trifurcula nov. spec., van Nieukerken in Kasy 1983: 5.
Type material. - Holotype &: Italy, M. Bondia, Villa
Faraldi, 3.5 km WNW Andora (Imperia), 550 m, UTM:
32T MP2769, 10.iv.1988, Dorycnium pentaphyllum,
stem-mines, e.l. 20.v.1988, EvN no. 88111, E. J. van Nieu-
kerken (RMNH). - Paratypes: 43 &, 13 9, 25 larvae. —
Austria: 1 @, Deutsch Altenburg, Pfaffenberg (Niede-
röst.), UTM: 33U XP43, 24.viii.1935, [Preissecker]
(NHMW) [paralectotype orientella Klimesch]; 8 8,
Hundsheimer Berg (Niederöst.), UTM: 33U XP43,
17.viii.1973, 28.vi.1976, 9.vi.1977, 3.vii.1977, 3.1x.1977,
22.v.1979, 12.ix.1979, E Kasy (NHMW, RMNH); 1 6,
Marchegg, Oberweiden (Niederöst.), UTM: 33U XP44,
6.1x.1931, [Preissecker] (NHMW) [ paralectotype orzen-
tella Klimesch]; 1 &, Mödling, Frauenstein (Niederöst.),
UTM: 33U WP92, 26.vi.1902, Preissecker (NHMW)
225
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
| paralectotype orzentella Klimesch |. - Czechoslovakia: 2
4, Hrädok n. V. (Slovakia), UTM: 33U YP19, 17.vi. 1988,
A. Lastivka; 2 8, Zádiel (Slovakia), UTM: 33U DU88,
7.vi. 1989, A. Lastúvka (coll. Laëtuvka, RMNH).- Greece:
1 8, Kardhamili (Messinias), gorge, UTM: 34S FF1084,
18.11.1990, Dorycnium hirsutum, stem-mines, el.
15.iv.1990, EvN no. 90063, E. J. van Nieukerken
(RMNH). - Italy: 1 8, Andora, Testico (Savona), UTM:
32T MP27, 2-5.vii.1983, B. À. Bengtsson (coll. Bengts-
son); 3 4, 2 9, Conna (Savona), 300 m, UTM: 32T MP27,
10.ix.1977, 1.1x.1978, 22.vi.1979, G. Baldizzone (GBA,
RMNH); 4 8,69, 1 larva, M. Bondia, Villa Faraldi, 3.5
km WNW Andora (Imperia), 550 m, UTM: 32T
MP2769, 10.iv.1988, Dorycnium pentaphyllum, stem-
mines, e.l. 19.v.1988-4.vi.1988, EvN no. 88111, E. J. van
Nieukerken (RMNH); 5 8, Poggio di Casasco, Val Cu-
rone (Al), 300 m, UTM: 32T NQ06, 06.vi.1987, G. Bal-
dizzone (GBA, RMNH). - Spain: 1 6, 1 9, 6 larvae, 2.5
km SW Beires, along road Beires-Fondón (Almería),
1000 m, UTM: 30S WF1495, 10.1.1988, Dorycnium pen-
taphyllum, stem-mines, e.l. 20.v.1988-26.vi.1988, EvN
no. 88017, van Nieukerken & Richter (RMNH); 1 4,
Collado de Falset (Cataluna), UTM: 30T CF15, 3.vii.1967,
E. Arenberger (LNKD); 1 4,29, 18 larvae, Sierra Blanca,
Refugio de Juanar, 3 km NW Ojén (Málaga), 840 m,
UTM: 30S UF3149, 15.1.1988, Dorycnium hirsutum,
stem-mines, e.l. 20.v.1988-8.vi.1988, EVN no. 88048, van
Nieukerken & Richter (RMNH); 2 9, Sierra de Marbella,
El Mirandor (Málaga), 700 m, UTM: 30S UF34,
21.vii.1987, E. Traugott-Olsen (ETO); 2 8, road Baza-
Benamaurel, 15 km from Baza (Granada), UTM: 30S
WG25, 16 and 17.vii.1987, Baldizzone & Traugott-Olsen
(GBA). - Soviet Union: 1 8, Parkovo, southcoast Crimea
(Krim), UTM: 36T WQ(southeast), 26.v.1984, Zaguljaev
(ZMAS). - Yugoslavia: 7 @, Krk, Draga Baska (Kroatia),
UTM: 33T VK78, 27.vii.1976, 28.viii.1978, 15.viii.1988,
G. Baldizzone (GBA, RMNH); 1 8, Krk, Skrpcici (Kroa-
tia), UTM: 33T VK68, 04.viii.1977, G. Baldizzone
(GBA).
Additional material. - Spain: mines, 3 km NE Mar-
bella, road to Ojén (Málaga), 200 m, UTM: 30S UF3244,
15.1.1988, Dorycnium hirsutum, EvN no. 88046, van
Nieukerken & Richter (RMNH).
Additional records. - Czechoslovakia: Cebovce (UTM:
34U CU63), Jablonov (34U DV83), Kiarov (34U CU83),
Hustopete (33U XQ22), Zajeti, larvae and adults, 1990,
leg. A. and Z. LaStuvka (Z. LaStuvka in litt.).
Description
Male (fig. 6). - Forewing length 2.3-3.0 mm (2.57
+ 0.18, 26), wingspan 4.9-6.5 mm. Head: frontal
tuft yellowish white to pale yellowish orange, collar
paler. Antenna with 37-43 (40.1 + 2.1, 22) seg-
ments; scape white, usually with some brown scales.
Forewing and thorax covered with fuscous tipped
scales, distal 4 to % of scales dark, edge darker, total
impression greyish brown, occasionally few white
scales exposed at dorsum; terminal cilia white
beyond indistinct cilia-line. Underside of forewing
with very large costal fold along basal third, com-
pletely covering a pocket with yellow androconial
scales of about one quarter wing width (figs. 20, 58).
Hindwing with pronounced humeral lobe, along
costa with long snow-white hair-scales, inserted on
underside, in rest fitting in forewing fold; imme-
226
diately behind frenulum a small group of closely set,
short dark fuscous special scales (figs. 21, 22, 58).
Female. - Forewing length 2.2-2.8 mm (2.52 +
0.19, 11) wingspan 4.9-6.1 mm. Antenna with 35-
38 (36.7 + 1.3, 10) segments.
Male genitalia (figs. 69-75) - Capsule length 340-
385 wm (363.6 + 14.5, 11). Vinculum anteriorly
truncate. Uncus with medial process distinctly wi-
dened terminally and truncate. Gnathos slightly or
not asymmetrical, central element narrowly spatu-
late, with a medial keel ventrally, margins often
serrate, additional spines may occur on ventral sur-
face, but specimens without any spines and smooth
margin do occur (figs. 69, 72, 73); anterior apo-
demes conspicuous. Valva length 260-290 um
(273.6 + 8.3, 11), almost triangular, with inward
curved tip; dorsally with a row of several spines
along continuation of lateral arm of transtilla,
transverse bar of transtilla slightly longer than sub-
lateral processes to twice as long. Aedeagus 335-385
um (360.0 + 17.3, 11) long, with ventral carina
fringed or bifid, aedeagal tube terminally spatulate,
dorsal lobe prominent at right side, with serrate
margin; vesica with two large spine-like cornuti,
one (155-200 um, 175.5 + 13.5, 10) basically joined
to a conical cornutus (50-90 um, 59 + 11.4, 10), the
other subterminally bent, partly hidden amongst
group of spine-like cornuti (not measurable); small
cornuti numerous. Juxta fig. 70.
Female genitalia (figs. 93, 99). - Terminalia dis-
tinctly pointed. T8 with 6-15 setae and some scales;
posterior margin truncate; anal papillae with 9-21
setae. Anterior apophyses laterally widened. Ductus
spermathecae with 4/2 to 5 coils. Signa 340-500 um
long, almost completely similar in length.
Final instar larva (fig. 104) - Yellow, elongate.
Headcapsule 375-425 um long, 1.01-1.11 (1.06 +
0.03, 6) X as long as wide (in one specimen 1.19 X).
Headcapsule distinctly longer than in subnitidella
and coronillae. Spinosity: prothorax ventrally with
spinose band posteriorly, meso- and metathorax
ventrally with a band along anterior margin and
few spines laterally. Abdominal segments 1-9 with
spines ventrally, not conspicuous; segment 10 with
small group of spines anterior of setae.
Diagnosis
Male immediately recognized by large costal fold,
hiding the yellow patch (N.B. this fold may be
difficult to see in well mounted specimens!), the
humeral lobe of the hindwing and the black scales
near the frenulum. Male genitalia in particular
characterized by gnathos with medial keel, but note
variability in gnathos! Female similar to other spe-
cies in the group, but easily separated by the pointed
postabdomen.
VAN NIEUKERKEN: Trifurcula subnitidella group
Figs. 69-75. Trifurcula josefklimeschi, male genitalia. - 69, Capsule, valva outlined, slide 2590; 70, Juxta, slide 2590; 71,
Valva, dorsal aspect, slide 2590; 72, 73, Gnathos, showing variation, respectively slides 3068 and 2744 (holotype); 74,
Aedeagus, ventral aspect, holotype, slide 2744; 75, Large cornuti, separately drawn, holotype. Scale 0.1 mm.
Biology
Hostplants. - Dorycnium hirsutum (L.) Ser. and
D. pentaphyllum Scop., both perennial herbs or
small shrubs, widely distributed in southern Eu-
rope, probably also on the other two European
Dorycnium species (D. rectum (L.) Ser. in DC. and
D. graecum (L.) Ser. in DC.).
Life history. - Egg usually deposited on the un-
derside of a leaflet, occasionally on the stem. When
in a leaflet, the larva starts with a short and narrow
linear mine (fig. 105), leading either in a straight
or slightly contorted course towards the petiole,
through which it enters the stem (fig. 105). The
larva then usually feeds downward first, in a rather
straight line, often doubling back once or twice,
before the larva quits the mine through a semicir-
cular slit. The whole course of the mine often stain-
ing reddish or brown, frass hardly visible externally,
larva visible as a yellow swelling in the stem. The
mine is frequently in the more terminal shoots, and
can relatively easily be found because of the stain-
ing. When full-grown larvae or empty mines are
collected, the leaflet with the early mine has often
already been fallen.
Larvae have been found in January, February and
early April. Adults emerged in mid April (from
227)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
© josefklimeschi
V victoris
Fig. 76. Distribution of Trifurcula victoris and T. josefklimeschi, mapped on 50 X 50 km UTM squares.
Greece), the second half of May and in June (indoor
rearing), but were caught from late May until the
12th of September, the largest numbers being taken
in June and from late August until early September.
This pattern might indicate a bivoltine cycle, but
this should be confirmed by finding larvae in
summer.
Distribution (fig. 76)
Widely distributed in southern Europe, from
southern Spain eastward to the Crimea, northwards
to eastern Austria and Slovakia. Not yet recorded,
but to be expected from Portugal, France, Hungary,
Rumania, Bulgaria and Turkey.
Remarks
Material of this species has long been known in
collections, but was incorrectly identified as Trifur-
cula orientella Klimesch. In fact, all four specimens
from Austria, listed amongst the type-material
by Klimesch (1953: 169) appeared to belong
to this species, which clearly differs from the spec-
imen which genitalia were illustrated by Klimesch
(1953: fig. 15) and is selected lectotype of orientella
below.
Etymology
A noun in genitive case, named in honour of Dr.
Josef Klimesch, who did much pioneer work on the
genus Trifurcula and on mediterranean Nepticuli-
dae in general.
228
8. Trifurcula iberica sp. n.
(figs. 29, 59, 77-82)
Type material. - Holotype @: Spain, Sierra Nevada,
road to Veleta (Granada), 1700 m, UTM: 30S VG60,
9.vii.1971, E. Arenberger (LNKD). - Paratype: 1 &:
Spain, Pyrenees, Prullans, 900 m, UTM: 31T CG99,
2.vii.1980, G. Derra (coll. Derra).
Description
Male. - Forewing length 2.7-3.2 mm, wingspan
6-7.2 mm. Head: frontal tuft ochreous yellow, collar
paler. Antenna with 37-40 segments; scape yellow-
ish white, with some brown scales. Forewing and
thorax covered with brown tipped scales, total im-
pression pale brown; terminal cilia white beyond
more or less distinct cilia-line. Underside of fore-
wing with basal patch of yellow androconial scales
extending to 4. Hindwing with white hair-scales
instead of costal bristles; along costa a row of short
but conspicuous black special scales, running from
4 to middle of hindwing (fig. 59).
Female unknown.
Male genitalia (figs. 77-82) - Capsule length 345-
355 um. Vinculum anteriorly truncate, rather nar-
row. Uncus with medial process pointed, some lat-
eral setae present. Gnathos almost symmetrical or
slightly asymmetrical, central element truncate to
rounded, posterior margin distinctly serrate; ante-
rior apodemes present. Valva length 275 um, nar-
row triangular, gradually narrowed towards tip,
dorsal excavation less than half length; transverse
bar of transtilla up to three times as long as sublat-
VAN NIEUKERKEN: Trifurcula subnitidella group
Figs. 77-82. Trifurcula iberica, male genitalia, slide 2741 (77-80) and 1928 (holotype: figs. 80, 82). - 77, Capsule, valva
outlined; 78, Juxta; 79, Valva, dorsal aspect; 80, Large cornuti, separately drawn; 81, Aedeagus, ventral aspect; 82,
Aedeagus, lateral aspect, position of spinelike cornutus shown. Scale 0.1 mm.
Diagnosis
The male of zberica is easily distinguished from
other species in the group by the long row of black
scales on the hindwing costa. The genitalia are
characterized by the almost symmetrical gnathos
with serrate margin, but resemble those of s¢/vzae,
see there.
Biology
Hostplant and immature stages unknown.
Adults taken in mountains, at medial (900 m) and
high altitude (1700 m), both in July.
Distribution (fig. 29)
Only known from two specimens from the Span-
ish Pyrenees and Sierra Nevada.
Etymology
A latin adjective, from Ibericus (= Hibericus),
meaning Spanish, referring to the single localities
in the Spanish mountains.
229
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
9. Trifurcula silviae sp. n.
(figs. 7, 29, 60, 83-88, 94, 100)
Type material. - Holotype 4: France, 1 km NW Ceillac,
+ 10 km S Chateau-Queyras (Htes Alpes), 1800 m, UTM:
32T LQ2449, 24.vii.1987, alpine meadow on S. slope,
netted at dusk, EVN no. 87139, van Nieukerken & Richter
(RMNH). - Paratypes: 6 6, 1 9. France: 1 8, data as
holotype (RMNH); 1 6, Chaner. [not traced, ?near
Digne], 22.viii.1903, Chrétien (MNHN); 2 @, Digne
(Alp. Ht. Prov.), UTM: 32T KP78, viii.1903, (MNHN);
1 4, Puy Vacher [near La Grave] (Htes Alpes), [2000 m],
UTM: 32T KQ88, 29.vi.1898, (MNHN); 1 6, Viens (Vau-
cluse), UTM: 31T GJ0664, 21.viii.1974, R. Buvat (coll.
Buvat); 1 9, same data, but 8.viii.1975 (RMNH).
Description
Male (fig. 7). — Forewing length 2.6-3.1 mm (2.80
+ 0.18, 6), wingspan 5.8-7.0 mm. Head: frontal tuft
pale yellow, mixed with white, collar yellowish
white. Antenna with 35-39 segments (36.5 + 1.5,
6) ; scape white. Forewing and thorax relatively
pale ochreous yellow, scales with ochreous brown
tips mixed with completely white scales; terminal
cilia white, cilia-line obsolete or absent. Underside
of forewing without patch of androconial scales,
narrow costal fold relatively well developed, edged
with row of brown scales (fig. 60). Hindwing al-
most white, costa of humeral lobe with row of short
dark brown scales, running to % (fig. 60).
Female. - Forewing length 2.6 mm, wingspan 5.8
mm. Antenna with 32 segments.
Male genitalia (figs. 83-88) - Capsule length 300-
330 um. Vinculum anteriorly truncate. Uncus with
medial process truncate, not widened, with several
lateral setae. Gnathos slightly or not asymmetrical,
central element narrow, posterior margin distinctly
serrate; anterior apodemes hardly visible. Valva
length 215-240 um, narrow triangular, tip short
pointed, slightly curved, transverse bar of transtilla
about 1.5 times as long as sublateral processes. Ae-
deagus 310-340 wm long, with ventral carina
fringed, aedeagal tube apically pointed, dorsal lobe
at right side prominent, margin serrate; vesica with
long pointed cornutus (140-210 um), basally joined
to conical cornutus (75 um), a long cornutus with
serrate tip hidden amongst group of long needle-
like cornuti, many small cornuti present. Juxta fig.
84.
Female genitalia (figs. 94, 100). - Terminal seg-
ments broadly rounded. T8 with about 6 setae and
few scales; anal papillae with 23-25 setae. Ductus
spermathecae with 4 coils. Signa 335-365 um long.
Diagnosis
Male differs from all other species in the subni-
tidella group by the absence of the yellow patch, it
is also paler than most species, except /uteola. T.
silviae male differs from other pale Trifurcula spe-
cies of similar size by the distinct rows of brown
230
scales along costal fold of forewing and along hindw-
ing costa. Male genitalia differ from similar iberica
by relatively longer sublateral processes and slightly
broader valva. Female differs from other species
described here by paler colour and blunt ovipositor.
Biology
Hostplant and immature stages unknown. The
holotype and one paratype were collected in an
alpine meadow on a steep southern slope, where
the Fabaceae Lotus corniculatus L., Anthyllis vulne-
raria L. and Onobrychis montana DC. are the most
likely candidates to be its host. The species appar-
ently occurs over a wide range of habitats, from
almost lowland mediterranean localities (Viens,
Digne), to high alpine country (Ceillac: 1800 m, Puy
Vacher: ca 2000 m.). Adults have been caught from
June to August.
Distribution (fig. 29)
Only known from a relatively small area in sou-
theastern France, in the Alps and pre-alps.
Remarks
The specimens in the Chrétien collection were
found amongst material of T. immundella, together
with a number of other misidentified species. The
single female is considered to belong to si/viae on
the basis of its occurrence with one male and no
other related species, the external characters and
the genitalia, which clearly belong to a species in the
subnitidella group.
Etymology
A noun in genitive case, named in honour of my
wife Silvia Richter, who not only collected both
holotype and one paratype, during an alpine moun-
tain hike, but contributed much to the collecting
during many joint field trips.
PHYLOGENY OF THE TRIFURCULA SUBNITIDELLA
GROUP
The phylogeny of the genus Trifurcula and its
three subgenera has been dealt with by Scoble
(1980) (with the present subgenera as genera) and
van Nieukerken (1986b). In these studies it has
been shown that the subgenus Glaucolepis Braun
(=Fedalmia Beirne) is the sistergroup of
[Levarchama Beirne + Trifurcula s.str.| and that
the latter two subgenera are sistergroups to each
other. Both the monophyly of the genus and subge-
neric clades has been amply demonstrated by apo-
morphies, to which little is to be added. Only one
more apomorphy for Levarchama has been menti-
oned by van Nieukerken & Johansson (1990): the
costal hair-pencil on the hindwing underside. The
apomorphy for Trifurcula s. str: uncus divided dor-
soventrally needs to be re-examined: in my opinion
VAN NIEUKERKEN: Trifurcula subnitidella group
Figs. 83-88. Trifurcula silviae, male genitalia. - 83, Capsule, valva outlined, holotype, slide 2742; 84, Juxta, slide 2604;
85, Valva, dorsal aspect, holotype; 86, Large cornuti, separately drawn, slide 2606; 87, Aedeagus, ventral aspect, slide
2606; 88, Aedeagus, lateral aspect, holotype. Scale 0.1 mm.
the ventral less sclerotized part, interpreted as part
of the uncus by Scoble (1980), might well be a
somewhat better sclerotized anal tube. This needs
to be confirmed by detailed examination of whole
mounts of complete genitalia, and might still be an
apomorphy for the subgenus. In any case, a new
interpretation of this character does in no way jeo-
pardize the monophyly of Trifurcula s. str.
The phylogeny within this subgenus has not been
discussed before, except the position of the two
South African species, discussed by Scoble (1980),
who could not solve the question whether they
together form the sistergroup of |Levarchama +
Trifurcula] or the sistergroup of Palaearctic Trifur-
cula s. str. I will come back to that question below,
and first concentrate on the Palaearctic species.
For an evaluation of the polarity of characters,
Levarchama has been used as outgroup. In this
study Trifurcula s. str. has been divided in two
species groups, which are believed to be monophy-
letic sistergroups. The phylogeny of the swbnitidella
group is presented in the cladogram in fig. 108.
Monophyly of the szbnitidella group is actually
based on one character only:
1. Male forewing with patch of yellowish androco-
nial scales on underside, near wingbase. Such andro-
conial scales are absent in Levarchama and in the
pallidella group, and not completely similar scales
are only found in a subgroup of Glaucolepis, so that
2511
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 89-94. Trifurcula spp., female postabdomen, dorsal aspect. - 89, T. puplesisi, slide 2763; 90, T. coronillae, slide 2750;
91, T. victoris, slide 2751; 92, T. subnitidella, slide 2746; 93, T. josefklimeschi, slide 2748; 95, T. silviae, slide 1818. Scale
0.1 mm.
232
VAN NIEUKERKEN: Trifurcula subnitidella group
95
100
Figs. 95-100. Trifurcula spp., female genitalia. - 95, T. puplesisi, slide 2763; 96, T. coronillae, slide 2750; 97, T. victoris,
slide 2752; 98, T. subnitidella, slide 2746; 99, T. josefklimeschi, slide 2748; 100, T. silviae, slide 1818.
255
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
101
103
102
Figs. 101-104. Trifurcula spp., final instar larva, headcapsule. - 101, T. coronillae, Spain, Istan; 102, T. victoris, Spain,
San José; 103, T. subnitidella, Netherlands, Kunrade; 104, T. josefklimeschi, Spain, Sierra Blanca.
the presence of these scales is regarded as an evo-
lutionary novelty. They are secondarily lost in s2/-
viae, which can be regarded to belong to the szbnz-
tidella group on the basis of other characters.
The pallidella group is believed to be monophy-
letic on the basis of at least the following apo-
morphy:
2. Hostplant belonging to the Genisteae (‘brooms’).
The hostplants of Levarchama and the subnitidella
group belong to other tribes of Fabaceae: Loteae
and Coronilleae, thus on the basis of the outgroup
rule, a hostplant belonging to Loteae or Coronilleae
is the plesiomorphous condition, and hence Genis-
teae apomorphous. No other Nepticulidae are
known to feed on brooms. Although only few spe-
cies have actually been reared, evidence is accumu-
lating that indeed all species are associated with
brooms.
234
3. A doubtful apomorphy is the form of the larger
cornuti: two are curved and one is spine-like in
virtually all species.
The current position of the first species of the
subnitidella group, T. austriaca, is tenuous. It is
included here solely on the basis of the yellow patch
and the general external resemblance to other spe-
cies in the group. The genitalia, in particular the
aedeagus, resemble more the pallidella group, al-
though they show some peculiarities, not noticed in
other species, such as the curved carinal lobes. If the
aedeagal characters (shape of cornuti, dorsal lobe)
belong to the groundplan of the subgenus, and aus-
triaca has no broom species as host, it still belongs
to the subnitidella group, and possibly is the sister-
species of all other species together. On the other
hand, if it is found to have a species of broom as
VAN NIEUKERKEN: Trifurcula subnitidella group
Figs. 105-107. Larval stem-mines of Trifurcula species. - 105, T. josefklimeschi, mine in Dorycnium hirsutum, Spain,
Sierra de Marbella; 106, T. coronillae, mines in Coronilla juncea, type locality; 107, T. subnitidella, mines in Lotus
corniculatus, Netherlands, Kunrade. Scale 1 cm.
295
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
AN
No S
d 9 9
9 ov A) A AV . è n ot
a 0 NI oÙ . LN x av R nv
CC o N N A N° N
Fig. 108. Cladogram of Trifurcula subnitidella group,
numbers refer to apomorphies, explained in text.
host, it is possibly misplaced here meaning that the
androconial patch has apparently evolved twice in-
dependently, or that it should be regarded as an
underlying apomorphy. Awaiting further informa-
tion of this species, austriaca is tentatively treated
here in the szbnitidella group.
The remaining eight species share the following
characters as apomorphies:
4. Aedeagus terminally narrowed, dorsal lobe well
developed, at right side or in middle; often with
serrate margin.
5. Pattern of the three large cornuti: one spine-like,
basally joined with a conical, and a long irregular
(often spine-like) cornutus hidden amongst the
group of needle-like cornuti.
The first branching in this clade has not been
resolved, and is provisionally presented as a tricho-
tomy. The species luteola and puplesisi resemble
each other closely, but the similarities seem to be
mere plesiomorphies, so that their phylogenetic
relationship cannot be demonstrated. The remain-
ing six species form a better defined monophyletic
entity on the basis of the apomorphy:
6. Gnathos highly atypical for Nepticulidae, with
keels and/or serrations, often highly asymmetrical.
The gnathos of the previous species resembles that
of the pallidella group, it is symmetrical and never
with keels, and thus forms the plesiomorphic con-
dition within the subgenus.
Also in the next clade the first branching is pres-
ented as a trichotomy, since the monophyly of
[coronillae + victoris] could not be demonstrated.
The monophyly of the final four species is based on
the following unique character:
7. Hindwing costa in male with characteristic black
or dark brown androconial scales
T. subnitidella and josefklimeschi are tentatively
regarded as sister-species on the basis of their sim-
ilarity and one possible apomorphy:
236
8. Black scales along hindwing costa confined to
small group close to frenulum.
Finally, T. zberica and silviae most likely are sis-
ter-species, also on the basis of a general similarity
in habitat, externals and genitalia of which the
following character might be regarded as apo-
morphy:
9. Gnathos without keels, but with serrated margin.
The phylogeny of the szbnitidella group is still
based on relatively few characters, more detail of
females, immature stages and hostplants would be
most welcome in order to refine the cladogram.
One final point to discuss here, is the phyloge-
netic position of the two South African species,
described by Scoble (1980).
Only males are known from these species, so that
we have no biological data to support our theories.
In my opinion both species definitely belong to
Trifurcula s. str. on the basis of the group of needle-
like cornuti, an apomorphy for the subgenus, and
the absence of a split uncus (in ventral view), a
character for Levarchama. However, the hair-pen-
cil of T. barbertonensis Scoble (fig. 12 in Scoble),
closely resembles that of Levarchama, which might
suggest another solution. Since no biological data
are available, and since it is unknown whether these
species have a patch of androconial scales on the
forewing underside, it is yet impossible to assign
them to one of the Palaearctic species groups. The
aedeagus as figured by Scoble (1980) seems to be
different from both species groups, so that it is still
possible that the South African species form the
sistergroup of the Palaearctic Trifurcula s. str.
APPENDIX
Trifurcula orientella Klimesch
Trifurcula orientella Klimesch, 1953: 168, 169, fig. 15.
Lectotype & (here designated): Yugoslavia, Dalmatia
mer., Umg. v. Gravosa, 15-31.v.1939, J. Klimesch, Ho-
lotypus’, genitalia slide Kl. 511 (coll. Klimesch)
[examined].
Trifurcula orientella was described on the basis of
six male specimens from Gravosa, Yugoslavia and
four male specimens from Austria. When I found
the austrian specimens in the Vienna museum, they
appeared to have genitalia differing from the figure,
presented by Klimesch, and actually appear to be-
long to the species here described as josefklimeschi.
Since Klimesch’s figure clearly has been the basis
for the identity of orientella, it seems most approp-
riate to select the specimen used for this figure as
lectotype. This is also the specimen labelled by
Klimesch ‘Holotypus’. Unfortunately, two other
paralectotypes from the type locality, examined by
me, also appear to belong to another and larger
species, either T. immundella Zeller or a closely
related species, which has been reared by me from
Calycotome. The remaining three paralectotypes
have not been examined by me.
T. orientella is a relatively small species, clearly
belonging to the pallidella group, which is known
from the Dalmatian coast, the isle of Krk (leg.
Baldizzone) and from Conna on the Italian Riviera.
From the latter locality I have reared this species
from stem-mines in the broom Genista germanica
IE,
ACKNOWLEDGEMENTS
The author is indebted for the loan of material
and information on distribution to the following
persons: G. Baldizzone (Asti, Italy), G. Bassi (To-
rino, Italy), B. A. Bengtsson (Löttorp, Sweden), R.
Buvat (Marseille, France), G. Derra (Bamberg, Ger-
many), A. M. Emmet (Saffron Walden, UK), R.
Gaedike (DEIC, Eberswalde, Germany), C. Gielis
(Lexmond, Netherlands), B. Gustafsson (NHRS,
Stockholm, Sweden), R. Johansson (Växjö,
Sweden), O. Karsholt (ZMUC, Copenhagen, Den-
mark), F. Kasy (NHMW, Vienna, Austria), S. Kerp-
pola (Helsinki, Finland), J. Klimesch (Linz, Aus-
tria), J. H. Kuchlein (Wageningen, Netherlands),
G.R. Langohr (Simpelveld, Netherlands), A. and Z.
Lastúvka (Brno, Czechoslovakia), G. Luquet
(MNHN, Paris, France), U. Parenti (Torino, Italy),
P. Passerin d’Entreves (Torino, Italy), R. Puplesis
(ZKVV, Vilnius, Lithuania), U. Roesler (LNKD,
Karlsruhe, Germany), H. Steuer (Bad Blankenburg,
Germany), I. Svensson ((Kristianstad, Sweden), E.
Traugott-Olsen (Marbella, Spain), K. Tuck
(BMNH, London, UK) and H. W. van der Wolf
(Nuenen, Netherlands). Ole Karsholt and Jan van
Tol are acknowledged for critical remarks on the
manuscript. Mrs. Ingrid Henneke assisted with
photographic work.
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Received: 26 September 1990.
Accepted: 1 October 1990.
ERIK J. van NIEUKERKEN
National Museum of Natural History, Leiden
STIGMELLA ROLANDI SP. N.: A WIDESPREAD
SOUTHERN EUROPEAN SPECIES ON ROSA
(LEPIDOPTERA: NEPTICULIDAE)
Nieukerken, E. J. van, 1990. Stigmella rolandi sp. n.: a widespread southern European
species on Rosa (Lepidoptera: Nepticulidae) — Tijdschrift voor Entomologie 133: 239-243,
figs. 1-10. [ISSN 0040-7496]. Published 14 December 1990.
Stigmella rolandi sp. n., belonging to the Stigmella sanguisorbae group, is described from
southern Europe. It has previously been misidentified as S. spinosissimae Waters, a western
European species. The species is characterized by a costal hair pencil on the male hindwing.
The distribution is mapped, and the biology described: the larva feeds on Rosa and Sangut-
sorba.
E. J. van Nieukerken, Rijksmuseum van Natuurlijke Historie, Postbus 9517, 2300 RA
Leiden, Netherlands.
Key-words. - Nepticulidae, leaf-miners, taxonomy, Palaearctic, Rosaceae.
The aim of this paper is to name a widespread
species of Stigmella, which has been known for
almost 40 years, but until recently was misidentified
as Stigmella spinosissimae (Waters). This misiden-
tification followed Klimesch (1951), who described
the genitalia and biology of the present species,
which he identified as S. spinosissimae, on the basis
of the same hostplant (Rosa pimpinellifolia L.) and
the description of external features by Waters
(1928). Study of type material of Nepticula spino-
sissimae Waters, however, showed that this is a
species in the anomalella species group, whereas
spinosissimae sensu Klimesch belongs to the san-
guisorbae species group (van Nieukerken 1986, Jo-
hansson & Nielsen 1990).
The European species of Stigmella Schrank are
relatively well known: Johansson & Nielsen (1990)
treated the 76 species of Northwest Europe in de-
tail. Only 20 additional species from southern Eu-
rope and the mediterranean region were listed by
van Nieukerken (1986). Most of these have been
satisfactorily described, including figures of male
genitalia, and in some cases female genitalia, by
Johansson (1971) and in papers by Klimesch (ref-
erences in Johansson & Nielsen 1990). To date only
about six more undescribed European species of the
genus are known in collections, and not many more
are expected to be found. Therefore, identification
of European Stzgmella species can be achieved with
a fairly high degree of certainty. In this light, des-
cription of a single widespread new species seems
justified and will facilitate identification. The other
undescribed species belong to different species
groups, mainly the ruficapitella and malella species
groups, and will be described in due time.
The Stigmella sanguisorbae group counts four
species (van Nieukerken 1986): of these, S. sangui-
sorbae (Wocke) and S. thuringiaca (Petry) have
been described and illustrated in detail by Johansson
& Nielsen (1990) and S. muricatella Klimesch in
the original description (Klimesch 1978). The
fourth species is (re)described and named below.
The methods and abbreviations are largely the
same as in the previous paper (van Nieukerken
1990), but genitalia measurements were taken at
400 X.
Stigmella rolandi sp. n.
(figs. 1-10)
[Nepticula spinosissimae Waters; Klimesch 1951: 4,
1958: 95, 1961: 751, Sz6cs 1955: 170, 1956: 390, 1963:
108, 1965: 59, 1973: 452, 1977: 92, 1978: 267, 1981:
213. Misidentifications]
Nepticula ‘spec. nov.’; Zimmermann 1944: 63.
Stigmella spinosissimae sensu Klimesch; van Nieukerken
1986: 9.
Type material. - Holotype @: Greece, Parnassós Oros,
5-6 km S. Polidhroson (Fokis), Abies-Pinus W., 1000-1200
m, 38.36N-22.33E [UTM: 34S FH37], 28.ix.1980, st. 59,
Rosa el. 25/29.iv.1981, VU no 80690KE, S. B. J. Menken
& E. J. van Nieukerken, Genitalia slide EvN 2780
(RMNH). - Paratypes: 81 4, 2 2. - Austria: 1 g, Dürn-
stein, UTM: 33U WP46, el 8.iv.1936, J. Klimesch
239
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 1,2. Stigmella rolandi, male. - 1 (top), habitus hol-
otype; 2 (bottom), Underside wings, showing special
scales and hair-pencil, Austria.
(RMNH). - Czechoslovakia: 1 8, Détkovice (Moravia),
UTM: 33U X057, 30.viii. 1988, el iii.1989, Rosa; 18,19,
Mikulov (Moravia), UTM: 33U XQ20, 29.ix.1989, el
11.1990, Sanguisorba minor; 2 6, SI. Kras-Zádiel, UTM:
Z. Laötúvka) — France: 1 4, La Penne-sur-l'Ouvèze
(Drôme), UTM: 31T FK70, 25-27.vi.1986 (coll. H. van
der Wolf); 4 8, Viens (Vaucluse), UTM: 31T GJ06,
22.vii. 1964, 24. viii.197 1, 8.viii.1975, 18.viii.1976, R. Buvat
(RMNH, coll. Buvat). — Italy: 1 @, Abruzzo, Sulmona
(l'Aquila), UTM: 34T EL88, 7.viii.1982 (coll. J. H. Kuch-
lein); 10 @, Monti Aurunci (Latina), 4 km NW Castel-
forte, UTM: 33T VF07, 400 m, 22-23.vi, 1.vii.1969; 2 &,
Monti Aurunci, 6 km N Itri, UTM: 33T UF77, 15.viii.1972
(coll. R. Johansson, RMNH); Sardegna, Mt. Istiddi, UTM:
31T NK12, 1.ix.1978 (coll. G. Derra); 1 6, Sardegna, Belvi
(Nuoro), UTM: 31T:NK12, 650 m, 20.vii.1984; 1 6, Sar-
degna, Fontana Raminosa (Nuoro), UTM: 31T NKI11,
900 m, 4.viii.1984; 3 8, Sardegna, Sarcidano (Nuoro),
UTM: 31T NK00, 720 m, 1.vii.1984 (all. leg. and coll. J.
H. Kuchlein). - Spain: 3 4, Cadalso de los Vidrios (Mad-
rid), UTM: 30T UK8062, 7.viii. 1986, E. J. van Nieukerken
(RMNH); 1 6,8 km E of Orcera (Jaen), 1150 m, UTM:
30S WH24, 19.vii.1986 (coll. Gielis); 5 &, San Roque
(Cadiz), UTM: 30S TF81, 26.vii.1986, C. Gielis (RMNH,
coll. Gielis); 2 &, Vega del Codorno (Cuenca), UTM: 30T
WK97, 1350 m, 23.vii. 1985 (coll. J. H. Kuchlein, RMNH).
— Yugoslavia: 1 g, Drenovo, Kavadarci (Macedonia),
UTM: 34T EL88, 1-10.vi.1957, FE Kasy (NHMW); 31 &,
1 9, Krk, Misucaynica, road Krk-Vrbnik (Kroatia), UTM:
33T VK78, 20.vii-11.viii, 1986-1988, G. Baldizzone
(RMNH, coll. Baldizzone); 3 &, Krk, Draga Baska (Kroa-
tia), UTM: 33T VK78, 15.viii.1988 (coll. Baldizzone). —
USSR, Ukraine: 1 8, Krim, Kara-Dag, 20 km W Feodosia,
240
UTM: 36T XQ78, 14.vii.1977, Reznik (ZMAS); 4 À, same
locality, 5-22.vii.1987, R. Puplesis (ZKVV, RMNH).
Description
Male (fig. 1).- Forewing length 1.6-2.1 mm (1.88
+ 0.11, 56), wingspan 3.7-4.7 mm. Head: frontal
tuft pale yellowish orange to ferruginous, collar
yellowish white. Antenna fuscous, with 25-29 seg-
ments (26.6 + 1.1, 44); scape yellowish white. Fore-
wing and thorax dark fuscous to fuscous black, scale
bases often paler greyish; terminal cilia dark grey,
occasionally separated by a more or less distinct cilia
line. Underside of forewing (fig. 2) with an elon-
gated andraconial patch, extending from base to %,
on costal side of fold, with fuscous grey special
scales, leaving a narrow furrow in middle; all scales
outwards oblique towards furrow. Hindwing grey,
first three to four costal bristles normal strong and
short, followed by group of more than 20 hairlike
costal bristles of Yo wing length, forming a hair-
pencil (fig. 3), which in rest is inserted in furrow in
forewing andraconial patch. Abdomen fuscous,
with distinct yellowish grey anal tufts.
Female. - Forewing length 1.8-1.9 mm (N=2),
wingspan 4.1-4.2 mm. Antenna with 20-22 seg-
ments. Forewing without special scales, hindwing
with costal bristles of normal length. No anal tufts.
Male genitalia (figs. 4-6) - Capsule length (from
tip of tegumen to central part of anterior margin of
vinculum) 163-210 um (188.4 + 12.5, 19). Vincu-
lum with anterior margin emarginate. Tegumen
hood-shaped. Uncus distinctly bilobed, lobes separ-
ate, each with some setae on prominent sockets.
Gnathos with posterior horns widely separate,
transverse bar anteriorly slightly protruding at
corners, forming indistinct anterior processes.
Valva length 148-180 um (164.2 + 8.4, 17), widest
beyond middle, suddenly tapering towards long
curved distal process; transverse bar of transtilla
long, sublateral processes small. Aedeagus 104-185
um (146.5 + 16.7, 17) long, tube broad, but slightly
variable in dimensions. Vesica with relatively few
small cornuti, some being a little larger.
Female genitalia (figs. 7, 8) — T8 with three
longitudinal bands of setae and scales; anal papillae
without setae. Posterior and anterior apophyses
long and narrow, approximately of same length.
Bursa globular, densely covered with pectinations,
no signum apparent. Accessory sac small, no retic-
ulate field visible. Ductus spermathecae without
distinct coils.
Diagnosis
The male can be recognized from all other uni-
formly coloured Stigmella species by the coastal
hairpencil and androconial scales on the forewing
underside. From above S. roland: resembles S. san-
guisorbae most, and females cannot be reliably se-
parated. S. thuringiaca has paler olive-brown to
grey-brown and more shining forewings, is some-
what larger, and usually has a darker head. Not
likely to be confused with species of the anomallela
group, that feed on the same hosts: S. anomalella
(Goeze) has distinct purplish wing tips, S. spinosis-
simae (Waters) has a fuscous head and bronze fore-
wings with copper reflections and S. centifoliella
(Zeller) has a postmedial fascia.
The male genitalia resemble also sanguisorbae
most, but roland: has much less cornuti. S thurin-
giaca has still more cornuti, plus pectinations on the
vesica and pectinate hairs on the dorsal face of the
valvae. Female genitalia with smaller accessory sac
than sangwisorbae or thuringiaca and without sig-
num. See illustrations in Johansson & Nielsen
(1990).
Biology
Hostplants. - Rosa spp., including R. pimpinel-
lifolia L. (= R. spinosissima L.) and Sanguisorba
minor Scop.
Mine (fig. 9). - A gallery mine. Egg deposited on
under- or upperside, often near a vein. Mine often
following the serrations of leaf-margin. Frass in
midline, leaving very narrow white margins in early
mine, but filling only about half mine width later.
Larva yellow. Mines are difficult or not to separate
from those of the anomalella group.
Life history. - Probably bivoltine. Larvae have
been found in late August and September, adults
reared from February to April. Early summer larvae
have not yet been found, but adults are found from
early June until early September, thus at least partly
from a probable second generation. Adults usually
taken at light.
Distribution (fig. 10)
Widespread in southern and southern central
Europe: eastern Austria, eastern Czechoslovakia,
southern France, Spain, Italy, Sardinia, Yugoslavia,
Greece and Soviet Union: Ukraine.
Literature records: Hungary: Budapest region
(Sz6cs 1955, 1956, 1963, 1981), Bakony mountains
VAN NIEUKERKEN: Stigmella rolandi sp. n.
Fig. 3. Stigmella rolandi,
male hindwing: costal bris-
tles and costal hair-pencil
(Italy: Monti Aurunci). Scale:
0.2 mm.
(Szöcs 1973), Matra mountains (Szöcs 1977) and
Pilis mountains (Szöcs 1978).
Etymology
A noun in genitive singular. Named in honour of
my friend Roland Johansson, specialist of Nepticu-
lidae, and outstanding painter of these moths.
ACKNOWLEDGEMENTS
The author would like to thank the following
persons for the loan and/or gift of material: G.
Baldizzone (Asti, Italy), R. Buvat (Marseille,
France), G. Derra (Bamberg, Germany), C. Gielis
(Lexmond, Netherlands), R. Johansson (Växjö,
Sweden), the late F Kasy (Wien, Austria), J. H.
Kuchlein (Wageningen, Netherlands), A. and Z.
Lastúvka (Brno, Czechoslovakia), R. Puplesis (Vil-
nius, Lithuania) and H. W. van der Wolf (Nuenen,
Netherlands). The author is particularly indebted
to Roland Johansson, for the continuous stream of
information and the many discussions about nepti-
culid taxonomy, which amongst others led to the
present paper.
REFERENCES
Johansson, R., 1971. Notes on the Nepticulidae (Lepidop-
tera) I. A revision of the Nepticula ruficapitella group.
— Entomologica scandinavica 2: 241-262.
Johansson, R. & E. S. Nielsen, 1990. Tribus Nepticulini.
- In: Johansson, R. et al. The Nepticulidae and Opos-
tegidae (Lepidoptera) of NW Europe. — Fauna ento-
mologica scandinavica 23: 111-238, pls.
Klimesch, J., 1951. Zur Kenntnis der Genitalmorphologie
einiger Nepticula-Arten (Lep., Nepticulidae). -
Zeitschrift der Wiener Entomologischen Gesellschaft
36: 4-9.
Klimesch, J., 1958. Beiträge zur Kenntnis der Lepidop-
teren-Fauna der Wachau in Niederösterreich (Microle-
pidoptera). - Zeitschrift der Wiener Entomologischen
Gesellschaft 43: 17-22, 43-44, 76-77, 91-97.
Klimesch, J., 1961. Ordnung Lepidoptera. I. Teil. Pyralid-
ina, Tortricina, Tineina, Eriocraniina und Micropte-
rygina. - In. H. Franz (ed.). Die Nordost-Alpen im
241
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 4-9. Stigmella rolandi, genitalia and leafmine. - 4, Capsule male genitalia, slide 2563 (Spain: Cadalso); 5, Valva,
inner aspect, slide 2563; 6, Aedeagus, holotype, slide 2780; 7, Female terminal segments, dorso-lateral view, slide 2783
(Yugoslavia: Krk); 8, Bursa copulatrix, detail showing enlarged pectinations, slide 2783; 9, leaf-mine on Rosa sp. from
type-locality, one of three mines from which holotype was reared. Scales: 0.1 mm (figs. 4-8), 5 mm (fig. 9); 4-6 and detail
of 8 on same scale.
242
VAN NIEUKERKEN: Stigmella rolandi sp. n.
Fig. 10. Distribution of Stigmella rolandi, mapped on 50 X 50 km UTM squares.
Spiegel ihrer Landtierwelt, 2: 481-789. Universitätsverlag
Wagner, Innsbruck.
Klimesch, J., 1978. Beitrag zur Kenntnis der Nepticuli-
denfauna von Anatolien und der Insel Rhodos (Lepi-
doptera, Nepticulidae). — Tijdschrift voor Entomolo-
gie 121: 239-278.
Nieukerken, E. J. van, 1986. A provisional phylogenetic
check-list of the western palaearctic Nepticulidae, with
data on hostplants (Lepidoptera). - Entomologica
scandinavica 17:1-27.
Nieukerken, E. J. van, 1990. The Trifurcula subnitidella
group (Lepidoptera: Nepticulidae): taxonomy, distri-
bution and biology. — Tijdschrift voor Entomologie
133: 000-000.
Szócs, J., 1955. A budapesti Mártonhegy lepke-faunája. —
Folia entomologica Hungarica, s. n. 8: 157-171.
Szócs, J., 1956. Magyarország Nepticulidái (Lepidopt.)
(Die in Ungarn vorkommenden Nepticula-Arten (Le-
pidopt.)) — Folia entomologica Hungarica, s. n. 9: 381-
394.
Szöcs, J., 1963. A lepkehernyoók természetes tápnövényei.
[Die natiirlichen Futterpflanzen der Schmetterlings-
raupen.]- Folia entomologica Hungarica, s. n. 16: 83-
120.
Szöcs, J., 1965. Microlepidoptera I, Nepticulidae. - Fauna
Hungarica 76: 48-104.
Szócs, J., 1973. Adatok a Bakony aknázómoly faunájához.
Angaben zur Minierfliegen-fauna [sic!] des Bakony-
Gebirges. - A Veszprém megyei múzeomok Közlemé-
nyei 12: 451-455.
Szócs, J., 1977. Adatok a Mátra-Hegység Aknázómoly-
faunájához. (Data to the mining moth fauna found in
the Mts. Matra.) - Folia Historia naturalis Musei Ma-
trensis 4: 91-99.
Szócs, J., 1978. Adatok a Pili-hegység Aknázómoly fauná-
jához. (Data to the mining moths fauna from the Pilis
Mts.) — Folia entomologica Hungarica, s. n. 31: 265-
Di,
Sz6cs, J., 1981. Angaben über minierenden Motten aus
Budapest und Umgebung. - Folia entomologica Hun-
garica, s. n. 42: 209-220.
Waters, E. G. R., 1928. A new Nepticula from North
Wales. - Entomologist's monthly Magazine 64: 105-
106.
Zimmermann, F., 1944. Zur Kenntnis der Verbreitung der
Nepticuliden in den Reichsgauen Wien und Niederdo-
nau (Lepidopt.). - Zeitschrift der Wiener Entomolo-
gischen Gesellschaft 29: 3-6, 60-64, 78-91, 107-122.
Received: 18 October 1990
Accepted: 18 October 1990
243
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
244
TILA M. PEREZ! & WARREN T. ATYEO?
Laboratorio de Acarologia, Universidad Nacional Autonoma de México
“Department of Entomology, University of Georgia
NEW TAXA OF FEATHER MITES (ACARINA,
PTEROLICHIDAE) FROM MEGAPODES (AVES,
MEGAPODIIDAE)
Pérez, T. M. & W. T. Atyeo, 1990. New taxa of feather mites (Acarina, Pterolichidae) from
Megapodes (Aves, Megapodiidae). - Tijdschrift voor Entomologie 133: 245-249, figs. 1-10,
table 1. [ISSN 0040-7496]. Published 14 December 1990.
Ascetolichus gen. n. (Pterolichoidea, Pterolichidae) is established for the type species
Pterolichus (Pseudalloptes) palmiger Trouessart and two new species, A. microthrix and
A. ruidus. All hosts are Megapodiidae.
Correspondence: W. T. Atyeo, Department of Entomology, University of Georgia, Athens,
Georgia 30602, U.S.A.
Key words. — Feather mites, Pterolichidae, Ascetolichus, Megapodiidae parasites.
In the late 1800s and early 1900s E. L. Trouessart,
often in collaboration with other authors, named
many taxa of feather mites taken primarily from
study skins in ornithological collections in Paris and
Angers. His collections included mites from three
species of Megapodiidae: ‘… les Talegalles
(Talegallus Cuvieri et Aepipodius Bruijnini) de
Nouvelle-Guinée’ and ‘... le Mégapode de Jobi
(Megapodius Jobiensis) de île Jobi (Nouvelle-Gui-
née). From these hosts, Trouessart (1887) and
Trouessart & Neumann (1888) described seven
mite species, however, the location of most of the
types is unknown. By examining megapode study
skins we have recovered the seven species of
Trouessart plus many new species.
From this rich and diverse acarofauna, we will
establish herein a new genus for one of seven
Trouessart species and two new species.
MATERIALS AND METHODS
We have examined 54 collections of feather mites
from 95 museum study skins of megapodes at the
American Museum of Natural History (see Atyeo
& Braasch 1966 for collecting technique). These
mites are from 13 of the 19 megapode species as
recognized by White & Bruce (1986). Each collec-
tion contains all mite specimens taken from an
individual skin, but the specimens may include only
a portion of the species known from the host spe-
cies (see Méjia-Gonzalez & Pérez 1988 for limita-
tions of the collecting technique). The megapode
species examined followed by the number of sam-
ples taken and the number of skins examined are
given in table 1. Names denoted by asterisks are
hosts from which species of the new genus were
collected.
All taxa being described, regardless of the sample
size, are believed to be from the Megapodiidae as we
have never encountered similar forms in any of our
extensive collections from the birds of the world.
Signatures for idiosomal setae follow the Grand-
jean system as modified by Griffiths et al. (1990).
Table 1. The number of feather mite collections (left
number) obtained from museum study skins examined
(right number) of megapodes. Asterisks indicate hosts of
Ascetolichus species.
Megapodius freycinet Gaimard, 1823 11 of 15
M. eremita Hartlaub, 1867 3 of 4
M. affinis A. B. Meyer, 1874
(= decollatus Oustalet, 1878) 1 of 3
(= jobiensis Oustalet, 1881) 3 of 5
M. laperouse Gaimard, 1823 9 of 14
M. pritchardii G. KR. Gray, 1864 0 of 10
M. (= Eulipoa) wallacei G. R. Gray, 1860 5 of 7
Leipoa ocellata ocellata Gould, 1840 1 of 1
L. ocellata rosinae Mathews, 1912 0 of 2
Alectura lathami J. E. Gray, 1831* 2 of 11
Talegalla cuvieri Lesson, 1828 3 of 3
T. jobiensis jobiensis A. E. Meyer, 1874* 4 of 5
T. fuscirostris Salvadori, 1877 4 of 5
Aepypodius arfakianus (Salvadori, 1877)* 3 of 4
A. bruijnii (Oustalet, 1880)* 2 of 2
Macrocephalon maleo S. Müller, 1846 3 of 4
245
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 1-2. Ascetolichus palmiger (Trouessart). Ventral and dorsal aspects of male. Setal signatures follow Griffiths et
al. (1990).
N 200 um
Figs. 3-4. Ascetolichus palmiger (Trouessart). Ventral and dorsal aspects of female. Setal signatures follow Griffichs et
al. (1990).
246
Measurements, in micrometres, are the mean +
standard error (when N>10), followed in paren-
theses by the observed limits and number of obser-
vations. The SEM micrographs were taken with a
Philips 505 using dehydrated specimens collected
from museum study skins.
Holotypes of new species will be deposited in the
American Museum of Natural History. Abbrevia-
tions for other type repositories and accession
numbers of bird and mite collections are: AMNH,
American Museum of Natural History, New York;
TRT, Trouessart Collection, Paris; and UGA, Uni-
versity of Georgia, Athens.
Ascetolichus Pérez & Atyeo, gen. n.
Diagnosis. - Small pterolichine mites with quad-
rate idiosoma and gnathosoma; well-developed
dorsal shields; posterolateral angles of propodo-
soma extended into points; all idiosomal and leg
setae present; setae vi expanded, minutely
branched; setae c2, cp large, branched; setae c3
minute, ventrolateral; cupules ia conspicuous; epi-
merites I free; legs subequal; legs III, IV ventral;
pretarsi symmetrical, dentate; tarsi I and II shorter
than corresponding tibiae; genua I, II, tibiae I, II
with dorsal crests or pebbled surfaces; bases of sole-
nidia 0, p respectively elevated above tibiae, tarsi;
genua I, II with setae mG bi- or trifurcate. Male
idiosoma with shallow terminal cleft on which is
inserted leaflike psl and branched hl; setae e2
ovate, fringed; adanal discs small, about 8 in diame-
ter, edentate; tarsus IV with subterminal claw. Fe-
male with idiosoma parallel-sided, more or less
truncated posteriorly; external spermduct present;
various terminal setae elaborated (as in fig. 4);
oviporus and associated structures anterior of seju-
gal furrow.
Type species. - Pterolichus (Pseudalloptes) pal-
miger Trouessatt.
Etymology. - Contraction of asketos (Gr., cur-
iously wrought, ornament) and Pterolichus, mascu-
line.
Ascetolichus palmiger (Trouessart) comb. n.
(figs. 1-6, 8)
Pterolichus (Pseudalloptes) palmiger Trouessart, 1887:
116-7; Canestrini & Kramer 1899: 61.
Males. — Dorsal shields with small pits unequally
distributed; genua I, II with ridges parallel to axis;
genua I, II with setae mG, cG coarsely branched;
setae cp long, bifurcate, unequal branches; measure-
ments: length, including gnathosoma 279 + 2 (262-
290, 19), width 170 = 1 (162-177, 19), sce:sce 86.5
+ 0.8 (78.4-92.1, 19), c2 length 46.2 + 0.7 (39.2-
49.0, 17), cp length 58.4 + 0.7 (50.9-63.7, 19), hl
length 8.3 + 0.2 (7.8-9.8, 18), ps1 19.6 + 0.3 (17.6-
PEREZ & ATYEO: Feather mites of Megapodes
21.6, 18), gnathosomal length 45.2 + 0.4 (43.1-49.0,
19), gnathosomal width 46.9 + 0.3 (45.1-49.0, 17).
Females. - Proterosoma as in male; dorsal hys-
terosoma with small pits, especially along margins;
measurements: length, including gnathosoma 435
(424-463, 7), external spermduct about 13.7; width
230 (220-254, 7), sce:sce 120.4 (113.7-129.4, 7), c2
length 59.1 (54.9-66.6, 6), c3 length 87.1 (82.3-92.1,
6), e2 length 51.9 (50.9-54.9, 6), f2 length 22.7
(19.6-23.5, 7), psl 14.8 (11.8-17.6, 7), ps2 23.3
(19.6-27.4, 5), gnathosomal length 65.8 (64.7-68.7,
7), gnathosomal width 69.2 (62.7-76.4, 7).
Type data. - From Aepypodius bruns: New
Guinea, | Waigeu Island], lectotype 6, paralectotype
Q (TRT slide 37 A 1). i
Material examined. — The type series (see Re-
marks) and other specimens from Aepypodius
bruijnü: Indonesia: West Irian: Waigeu Island, 15
6, 4 9, circa 1894, A. A. Bruijn (AMNH 539411,
UGA 5339; AMNH 539410, UGA 5340). From A.
arfakianus; Indonesia: West Irian: Idenburg River,
6 km SW Bernhard Camp (39° 15’E, 3°30'S), 1 6,
3 9, 21 February 1939, R. Archbold (AMNH
338429, UGA 5336); Snow Mountains, 3 @ , 1 9, 24
September 1910, A. S. Meek (AMNH 539393, UGA
5337). From Talegallus cuvieri: New Guinea: no
other data, 2 Q (TRT slides nos. 37 A 2, 22 H 16).
Remarks. - The number of observations for setae
and gnathosomal measurements are small as many
setae are lacking from the specimens and many
gnathosomata have been deformed during speci-
men preparation. The measurements for the gnath-
osoma are included in this description only; they are
similar for the other species for which there are few
specimens.
Trouessart (1887) stated that the hosts for this
species were T. cuvieri and A. bruijnii, New Guinea.
Our collections from A. bruijnii are conspecific with
Trouessart’s specimens from the listed hosts (see
above). We assume that Trouessart’s two females
from T. cuvieri represent accidental associations.
Ascetolichus species
(fig. 7)
One of the males used for SEM micrographs is an
undescribed species (fig. 7) from the study skin of
Aepypodius arfakianus from the Bernhard Camp
(listed above). All males from the Aepypodius col-
lections have been mounted and additional speci-
mens were not found. Although the host associa-
tion must be tentative until additional specimens
are collected, it is possible that two species of Asce-
tolichus could occur on A. arfakianus. Two or three
congeners on a single host, with each congener
occuping a different microhabitat has been well
documented (Atyeo & Pérez 1988, Méjia-González
& Pérez 1988).
The species is related to Ascetolichus palmiger as
247
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
sog
Figs. 5-8. SEMs of dorsal aspects of Ascetolichus species. — 5, Ascetolichus palmiger, male: arrow = setae e2; 6, A.
palmiger, male propodosoma: upper arrow = setae cG, lower arrow = seta mG, genual ridges between setae; 7,
Ascetolichus species, male: arrow = setae psl; 8, A. palmiger, female. Scale bars: 5, 7,8 = 100 um; 6 = 50 um.
200 um
(3)
2
oe 099
Dos 09
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O
5
06
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0)
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OL
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ar
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oc
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Figs. 9-10. Dorsal aspects of Ascetolichus species. - 9, A. rwidus sp. n., female; 10, A. microthrix sp. n., male. G = position
of male genital organ.
248
shown by the ridges on the anterior legs, but the
dorsal ornamentation is different, setae c2 and cp
are smaller, and setae psl are bifurcated.
Ascetolichus ruidus Pérez & Atyeo, sp. n.
(fig. 9)
Female (holotype). - Dorsal idiosoma with large,
deep pits; genua I, II, femur II with antaxial surfaces
pebbled; tibiae I, II with setae mG, cG thin, bi- or
trifurcate; setae cp long, bifurcate, unequal
branches; measurements: length, including gnath-
osoma 424, width 197, external spermduct 43.1,
sce:sce 11.7, c2 66:6, cp 84.3, e2 3912, f2 19.6, ps]
DB 1982 197%
Male. - Unknown.
Type data. - The holotype female was collected
from Alectura lathami: Australia: Queensland: (?)
Barron River, 19 July 1884, G. M. Mathews
(AMNH 539322, UGA 5324). The holotype is dep-
osited in AMNH.
Etymology. - From rwidus (Gr., rough) to refer
to the rugose tegument of legs I and II.
Remarks - We have only two collections from A.
lathami and in one, the unique holotype was found.
Until validated by additional material, the host as-
sociation is considered tentative.
Ascetolichus microthrix Pérez and Atyeo, sp. n.
(fig. 10)
Males (N=2). - Dorsal idiosoma with large pits;
legs I, II without pebbled tegument or ridges paral-
lel to axis; genua I, II with setae mG simple or with
small basal branch; setae cp bifurcate, with few
branches; measurements: length, including gnath-
osoma 280 (278-282), width 164 (162-166), sce:sce
95.1 (94.1-96.0), c2 length 29.4 (27.4-31.4), cp
length 53.9 (49.0-58.8), hl length 11.8 (11.8), psl
27.9 (27.4-28.4).
Female (N=1). - Proterosoma and dorsal idio-
soma similar to male; measurements: length, in-
cluding gnathosoma 416, width 208, sce:sce 113.7,
external spermduct 29.4, c2 31.4, cp 74.5, e2 23.5,
PAG ps 137, \ps2 bles:
Type data. - From Talegalla j. jobiensis: Indone-
sia: West Irian: Humboldt Bay, Hollandia, holotype
6, 1 ©, 1 Q paratypes, 5 August 1928, E. Mayr
(AMNH 539377, UGA 5332). The holotype is dep-
osited in AMNH, paratypes in UGA.
PEREZ & ATYEO: Feather mites of Megapodes
Etymology. - From mikros (Gr., small) and thrix
(Gr. seta) to refer to the small setae c2.
Remarks. - The two males in this collection had
setae e2 missing; they have been represented with
dotted lines as structures similar to those of A.
palmiger.
The females of the two new species, A. rwidus and
A. microthrix differ from those of A. palmiger by
having long external spermducts. The female of A.
ruidus differs from A. microthrix in having the pits
of the hysterosomal shield approximately equal in
size, a longer spermduct, and longer setae (espe-
cially c2, e2, ps1, compare measurements).
ACKNOWLEDGEMENT
This research was supported by the National
Science Foundation (BSR 89-08301).
REFERENCES
Atyeo, W. T. & N. L. Braasch, 1966. The feather mite
genus Proctophyllodes (Sarcoptiformes: Proctophyllo-
didae). - Bulletin of the University of Nebraska State
Museum 5: 1-354.
Atyeo, W. T. & T. M. Pérez, 1988. Species in the genus
Rhytidelasma Gaud (Acarina: Pterolichidae) from the
Green Conure, Aratinga holochlora (Sclater) (Aves:
Psittacidae). — Systematic Parasitology 11: 85-96.
Canestrini, G. & P. Kramer, 1899. Demodicidae und Sar-
coptidae. - Das Tierreich 7: 1-193.
Griffiths, D. A., W. T. Atyeo, R. A. Norton & C. A. Lynch,
1990. The idiosomal chaetotaxy of astigmatid mites. —
Journal of Zoology, London 220: 1-32.
Méjia-González, E. & T. M. Pérez, 1988. Three new spe-
cies of Fainalges Gaud and Berla (Analgoidea: Xolal-
gidae) with descriptions of their developmental series.
— Acarologia 29: 73-86.
Trouessart, E. L. (1886) 1887. Diagnoses d'espèces nou-
velles de Sarcoptides plumicoles (Analgesinae). — Bul-
letin de la Société d'études scientifiques d'Angers 15:
85-156.
Trouessart, E. L. & G. Neumann, 1888. Diagnoses d’espe-
ces nouvelles de Sarcoptides plumicoles (Analgesinae).
- Bulletin scientifique de la France et de la Belgique 19:
325-380.
White, C. M. N. & M. D. Bruce, 1986. The birds of Wal-
lacea. - British Ornithologists’ Union Check-list No. 7,
524 p.
Received: 1 May 1990
Revised version accepted: 30 July 1990
LS)
Se
Ne)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
250
D. L. J. QUICKE! & C. van ACHTERBERG?
"Department of Animal and Plant Sciences, University of Sheffield
*Nationaal Natuurbistorisch Museum, Leiden
THE TYPE SPECIMENS OF ENDERLEIN'S
BRACONINAE (HYMENOPTERA:
BRACONIDAE) HOUSED IN WARSAW
Quicke, D. L. J. & C. van Achterberg, 1990. The type specimens of Enderlein's Braconinae
(Hymenoptera: Braconidae) housed in Warsaw. — Tijdschrift voor Entomologie 133: 251-
264. [ISSN 0040-7496]. Published 14 December 1990.
The type specimens of 147 species of Braconinae described by Enderlein in the collection
of the Polish Academy of Science, Institute of Zoology, Warsaw have been examined. Eighty-
one new combinations are reported, the systematic positions of 39 species are confirmed
and lectotypes are designated for 62 species. The genera Diolcia Enderlein and Monolcia
Enderlein are synonymized with Campyloneurus Szépligeti; Diamblomerina Enderlein is
synonymized with Nedinoschiza Cameron; Diamblomera Enderlein is removed from syn-
onymy with Odontoscapus Kriechbaumer and provisionally treated as a valid genus. Cya-
nopterus solox Enderlein is treated as a new junior synonym of Exurobracon triplagiata
Cameron.
Correspondence: D. L. J. Quicke, Department of Animal and Plant Sciences, University of
Sheffield, Sheffield, U.K.
Key words. - Hymenoptera, Braconidae, Braconinae, Enderlein, reclassification, lectotypes.
Günter Enderlein (1905a, 1905b, 1920) described
a large number of new taxa of Braconidae including
153 new species and 17 new genera of Braconinae,
the vast majority being dealt with in the last of his
papers. The primary types of 147 taxa are listed in
this paper because they are deposited in the Instytut
Zoologii of the Polish Academy of Sciences in War-
saw (PAN). Enderlein worked at the Museum fiir
Naturkunde, Stettin (now Szczecin in north-west
Poland) between the years 1906 and 1919. The
principally tropical Braconidae dealt with in his
papers were mainly from the collection of the
former curator of the Stettin Museum, Dr. H.
Dohrn and most of them were collected in Sumatra
by Mr. M. Ude. The specimens in the collection
formerly belonging to the Museum fiir Naturkunde
in Berlin are duplicates retained by Enderlein. As a
consequence of the Second World War, the collec-
tion finally ended up in the Polish Academy of
Sciences. It is now housed at a field station/ museum
belonging to the PAN at Lomna, approximately 30
km north-west of Warsaw, and the main part of the
material is in very good condition.
Although it had been possible to study some of
this material in the past by means of small postal
loans, damage that accrued to a considerable part of
such loaned material due to bad postal handling has
meant that Dr Eusgeniusz Kierych, the curator of
entomology at the PAN, was reluctant to continue
such loans. Thus in order to examine the entirity of
the Enderlein collection so that the identities of
many of his species could be re-assessed, the authors
have recently visited the PAN facility at Lomna and
the findings are reported below.
Enderlein’s species concept seems to have been
reasonable, although he described the same species
on several occasions twice in the same paper; only
rarely are his type series heterogeneous. However,
his generic concept was, judging from his work on
the Braconinae, very poor indeed and he would
often describe two new genera based on one or two
closely related species. For example, his genera An-
tidiolcus and Plagiozina (both of which have now
been synonymized with Nesaulax Roman; Quicke
1984a) are based on what are in fact two sibling
species. Similarly, his genera Diolcia and Monolcia
are based on another extremely closely related pair
of species, both of which are transferred below to
Campyloneurus Szépligeti. On other occasions he
split the members of a single genus between as
many as four genera, for example, species of Hemz-
bracon Szépligeti were described under Ipobracon
Thomson, Antiolcus Enderlein and Udamolx En-
derlein as well as in Hemibracon itself.
Enderlein’s inconsistencies and the errors in the
generic level classification of his braconine species
251]
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 1. Overview of the current taxonomic status of the Braconinae described by G. Enderlein and deposited in PAN.
Current genus
Antiolcia Enderlein
Archibracon Saussure
Bathyaulax Szépligeti
Blastomorpha Szépligeti
Campyloneurus Szépligeti
Chaoilta Cameron
Chelonogastra Ashmead
Compsobracon Ashmead
Cordibracon van Achterb.
Craspedolcus Enderlein
Cratobracon Szépligeti
Curriea Ashmead
Diamblomera Enderlein
Digonogastra Viereck
Species
mitelligera Enderlein, 1920: 116-117
atricauda Enderlein, 1920: 97
gutta Enderlein, 1920: 97-98
flavipera Enderlein, 1920: 126-127
gutta Enderlein, 1920: 122-123
laeviventris Enderlein, 1920: 126
latiangulata Enderlein, 1920: 125-126
nigriceps Enderlein, 1920: 125
pectinatus Enderlein, 1920: 123
rugiventris Enderlein, 1920: 124-125
celebensis Enderlein, 1920: 55-56
angulosus Enderlein, 1920: 115
basalis Enderlein, 1920: 105-106
bicarinatus Enderlein, 1920: 112
cingulicauda Enderlein, 1920: 105
flavicosta Enderlein, 1920: 115-116
impressimargo Enderlein, 1920: 109
latispeculum Enderlein, 1920: 103
limbaticauda Enderlein, 1920: 112
marginiventris Enderlein, 1920: 109-
110
nigricosta Enderlein, 1920: 108
punctativentris Enderlein, 1920: 110
reticulatus Enderlein, 1920: 107
serenans Enderlein, 1920: 104-105
speculiger Enderlein, 1920: 109
tibialis Enderlein, 1920: 108
tricarinatus Enderlein, 1920: 113
trispeculatus Enderlein, 1920: 107-108
carinicornis Enderlein, 1920: 55
compta Enderlein, 1920: 56
disciventris Enderlein, 1920: 103
thodeanus Enderlein, 1920: 57
laqueatus Enderlein, 1920: 120
fraternus Enderlein, 1920: 92
maculicosta Enderlein, 1920: 71-72
obscuriventris Enderlein, 1920: 93
trisulcatus Enderlein, 1920: 92
gibber Enderlein, 1920: 65-66
grata Enderlein, 1920: 117
gratiosus Enderlein, 1920: 53
guttifer Enderlein, 1920: 53
acuticellus Enderlein, 1920: 100-101
grisescens Enderlein, 1920: 101
acuticampa Enderlein, 1920: 86
alboniger Enderlein, 1920: 78
aterimma Enderlein, 1920: 98
aureopilosa Enderlein, 1920: 80
biareata Enderlein, 1920: 92
bicristata Enderlein, 1920: 88
bicuneata Enderlein, 1920: 87-88
bicunea Enderlein, 1920: 83-84
brevicapula Enderlein, 1920: 91
brevicunea Enderlein, 1920: 78-79
camerom Enderlein, 1920: 82
chilensis Enderlein, 1920: 58
columbiana Enderlein, 1920: 82
crenulata Enderlein, 1920: 79
digitata Enderlein, 1920: 84
duploareata Enderlein, 1920: 82-83
ecuadorensis Enderlein, 1920: 85
fenestrata Enderlein, 1920: 89
252
Original genus
Antiolcia
Udamolx
Udamolx
Euryacria
Gontobracon
Bathyaulax
Bathyaulax
Bathyaulax
Gontobracon
Bathyaulax
Platybracon
Monolcia
Campyloneurus
Diolcia
Campyloneurus
Monolcia
Campyloneurus
Campyloneurus
Diolcia
Campyloneurus
Campyloneurus
Campyloneurus
Campyloneurus
Campyloneurus
Campyloneurus
Campyloneurus
Monolcia
Campyloneurus
Platybracon
Platybracon
Campyloneurus
Cyanopterus
Udamolcia
Craspedolcus
Ipobracon
Craspedolcus
Craspedolcus
Merinotus
Antiolcia
Aphrastobracon
Aphrastobracon
Diamblomera
Diamblomera
Ipobracon
Ipobracon
Udamolx
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Cyanopterus
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Type-locality
Cameroons: Barombi
S. Africa: Natal
S. Africa: Natal
Malagasy: Amber Mountains
S. Africa: Natal
Malagasy: Amber Mountains
Malagasy: Tamatave
S. Africa: Transvaal: Zout-
pansberg
S. Africa: Transvaal: Zout-
pansberg
Tanzania
Indonesia: N. Sulawesi
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Taiwan: Taihorin
Indonesia: Java: Sukabumi
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Indonesia: Java: Sukabumi
Sukaranda
Sukaranda
Sukaranda
Sumatra:
Sumatra:
Sumatra:
Indonesia:
Indonesia:
Indonesia:
Ceylon
Indonesia:
Indonesia:
Indonesia:
Indonesia:
Indonesia:
Indonesia:
Indonesia: Sumatra: Liangagas
Indonesia: Sumatra: Sukaranda
Brazil: Santa Catharina
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Indonesia: Java: Sukabumi
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Tanzania: Nyembe Bulungwa
Bioka (= Fernando Poo)
Cameroons: Barombi
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Colombia: Rio Magdalena
Colombia
Brazil
Brazil: Obodos
Peru
Colombia
Brazil: Obidos
Ecuador: Curaray
Guyana: Demerara
Peru: Chanchamayo: Rio Toro
Mexico: Chiapas
Chile
Colombia
Brazil (? Guyana): Demerara
Brazil: Santa Catharina
Brazil: Santa Catharina
Ecuador: Balzapampa
Brazil: Pebas
Sukaranda
Sukaranda
Sukaranda
Sukaranda
Sukaranda
Sukaranda
Sumatra:
Sumatra:
Sumatra:
Sumatra:
Sumatra:
Sumatra:
Table 1. (continued).
Current genus
Ectemnoplax Enderlein
Euwrobracon Ashmead
Gammabracon Quicke
Gronaulax Cameron
Hemibracon Szépligeti
Hybogaster Szépligeti
Iphiaulax Foerster
Ischnobracon Baltazar
Isomecus Kriechbaumer
Megabracon Szépligeti
Monocoila Roman
Nedinoschiza Cameron
Nesaulax Roman
Pachybracon Cameron,
-group !)
Physaraia Shenefelt
Plaxopsis van Achterberg
Plesiobracon Cameron
Rbadinobracon Szépligeti
Rhamnura Enderlein
Serraulax Quicke
Shelfordia Cameron
Spathulibracon Quicke
Stenobracon Szépligeti
Syntomernus Enderlein
Trigastrotheca Cameron
Trispinaria Quicke
QUICKE & VAN ACHTERBERG: Enderlein’s Braconinae
Species
flavicaligata Enderlein, 1920: 80
latecrenulata Enderlein, 1920: 81-82
latefasciata Enderlein, 1920: 90
laticampa Enderlein, 1920: 90-91
laticunea Enderlein, 1920: 79
longicapula Enderlein, 1920: 91
macella Enderlein, 1920: 86
mediofusca Enderlein, 1920: 88-89
nigripecta Enderlein, 1920: 83
obtusicampa Enderlein, 1920: 85-86
ochripes Enderlein, 1920: 86-87
penniseta Enderlein, 1920: 87
rectivena Enderlein, 1920: 81
sigillata Enderlein, 1920: 79
speculata Enderlein, 1920: 89-90
subfracta Enderlein, 1920: 96-97
tuberculata Enderlein, 1920: 84-85
peruliventris Enderlein, 1920: 110-111
merinotoides Enderlein, 1920: 128-129
solox Enderlein, 1920: 58
nigriseta Enderlein, 1920: 68-69
laticampus Enderlein, 1920: 69
apicalis Enderlein, 1920: 96
catharinensis Enderlein, 1920: 98-99
crassecrenulatus Enderlein, 1920: 81
nigripalpalis Enderlein, 1920: 90
politus Enderlein, 1920: 99
vaucristatus Enderlein, 1920: 99-100
w-impressus Enderlein, 1920: 63
crista Enderlein, 1920: 75-76
breviseta Enderlein, 1920: 60
sautert Enderlein, 1920: 128
udet Enderlein, 1920: 127-128
rhyssides Enderlein, 1920: 61-62
quadrirugulosus Enderlein, 1920: 94
filiseta Enderlein, 1920: 57-58
lurida Enderlein, 1920: 111
speciosus Enderlein, 1920: 121-122
excisus Enderlein, 1920: 95-96
gracilis Enderlein, 1920: 67-68
pravivena Enderlein, 1920: 94-95
albescens Enderlein, 1920: 117-118
albicans Enderlein, 1920: 102-103
apicalis Enderlein, 1920: 114
gibbiventris Enderlein, 1920: 104
insolita Enderlein, 1920: 118
sumatrana Enderlein, 1905b: 236
magnificus Enderlein, 1920: 77-78
cincticauda Enderlein, 1920: 104
ruficauda Enderlein, 1920: 71
capillicauda Enderlein, 1905a: 196
filicauda Enderlein, 1905a: 195
denticornis Enderlein, 1920: 123
bispeculum Enderlein, 1920: 69
criniseta Enderlein, 1920: 68
flagriseta Enderlein, 1920: 64-65
ingentiseta Enderlein, 1920: 70-71
quadricarinata Enderlein, 1920: 66-67
rimicunea Enderlein, 1920: 67
rugiventris Enderlein, 1920: 77
elegantulus Enderlein, 1920: 62-63
pusillus Enderlein, 1920: 121
tridentata Enderlein, 1920: 60-61
sannio Enderlein, 1920: 54
Original genus
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Ipobracon
Antiolcus
Ipobracon
Ectemnoplax
Bracon
Cyanopterus
Merinotus
Merinotus
Antiolcus
Udamolx
Ipobracon
Ipobracon
Udamolx
Udamolx
Hemibracon
Ipobracon
Megagonia
Iphiaulax
Iphiaulax
Rhadinobracon
Craspedolcus
Cyanopterus
Ectemnoplax
Diamblomerina
Antidiolcus
Merinotus
Plagiozina
Udamolcia
Campyloneurus
Monolcia
Campyloneurus
Udamolcia
Gastrotheca
Ipobracon
Campyloneurus
Meronotus
Rhamnura
Rhamnura
Goniobracon
Merinotus
Merinotus
Merinotus
Merinotus
Merinotus
Merinotus
Ipobracon
Hemibracon
Syntomernus
Odontopygia
Pseudospinaria
Type-locality
Guyana: Demarara
Brazil: Santa Catharina
Brazil: Obidos
Guyana: Demerara
Peru: Chanchamayo: Rio Toro
Guyana: Demerara
Peru: Chanchamayo
Brazil: Obidos
Argentine: Mendoza
Colombia
Ecuador: Guayaquil
Brazil: Obidos
Brazil: Santa Catharina
Peru
Colombia: Rio Magdalena
Brazil: Obidos
Peru: Chanchamayo: Rio Toro
Taiwan: Takao
Indonesia: Sumatra: Sukaranda
India: Sikkim
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Costa Rica
Brazil: Santa Catharina
Brazil: Santa Catharina
Ecuador: Balzapamba
Peru: Chanchamayo
Guyana: Demerara
Brazil: Sao Paulo de Olivenca
Indonesia: Sumatra: Sukaranda
Indonesia: Sumba
Taiwan: Takao
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Ecuador: Bucay
Peru: Chanchamayo: Rio Toro
S. Africa
Indonesia: Sumatra:
Indonesia: Sumatra:
Indonesia: Sumatra:
Indonesia: Sumatra:
Indonesia: Sumatra:
Indonesia: Sumatra:
Indonesia: Sumatra:
India
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Tanzania: Usambara: Bulwa
Indonesia: Sumatra: Liangagas
Namibia
Togo: Bismarckburg
Cameroons: Joh.-Albrechtshöhe
Cameroons: Kribi
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Indonesia: Java: Sukabumi
Indonesia: Sumatra: Sukaranda
Taiwan: Takao
Indonesia: Sumatra: Sukaranda
Indonesia: Sumatra: Sukaranda
Sukaranda
Sukaranda
Sukaranda
Sukaranda
Sukaranda
Sukaranda
Sukaranda
255
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Table 1. (continued).
Current genus
Undabracon Quicke
Zaglyptogastra Ashmead
Species
jucundus Enderlein, 1920: 119
griseiseta Enderlein, 1920: 59
helvimacula Enderlein, 1920: 123-124
plumiseta Enderlein, 1920: 59-60
Original genus Type-locality
Udamolcia Indonesia: Sumatra: Sukaranda
Megagonia Indonesia: Sumatra: Sukaranda
Goniobracon Ethiopia: Eritrea: Asmara
Megagonia Indonesia: Sumatra: Sukaranda
Unplaced species (these taxa will be treated in a forthcoming paper by the junior author)
albimanus Enderlein, 1920: 118-119
angustisulca Enderlein, 1920: 73-74
basispeculum Enderlein, 1920: 73
consimilis Enderlein, 1920: 113-114
denticornis Enderlein, 1920: 120-121
flavimarginatus Enderlein, 1920: 114
gracilis Enderlein, 1920: 118
laevibasis Enderlein, 1920: 75
laeviventris Enderlein, 1920: 76-77
latisulca Enderlein, 1920: 74
maculistigma Enderlein, 1920: 107
parvispeculum Enderlein, 1920: 73
serenimanus Enderlein, 1920: 114-115
sexrugosus Enderlein, 1920: 72-73
strigidorsum Enderlein, 1920: 93
trirugosus Enderlein, 1920: 72
trispeculum Enderlein, 1920: 70
undicuneus Enderlein, 1920: 106-107
Udamolcia Indonesia: Sumatra: Sukaranda
Ipobracon Indonesia: Sumatra: Sukaranda
Ipobracon Indonesia: Sumatra: Sukaranda
Monolcia Indonesia: Sumatra: Sukaranda
Udamolcia Indonesia: Sumatra: Sukaranda
Monolcia Indonesia: Java: Sukabumi
Udamolcia Indonesia: Sumatra: Sukaranda
Ipobracon Indonesia: Sumatra: Sukaranda
Ipobracon Indonesia: Sumatra: Sukaranda
Ipobracon Indonesia: Sumatra: Sukaranda
Campyloneurus Indonesia: Sumatra: Sukaranda
Ipobracon Indonesia: Sumatra: Sukaranda
Monolcia Indonesia: Sumatra: Sukaranda
Ipobracon Indonesia: Sumatra: Sukaranda
Craspedolcus Indonesia: Sumatra: Sukaranda
Ipobracon Indonesia: Sumatra: Sukaranda
Merinotus Indonesia: Sumatra: Sukaranda
Campyloneurus Indonesia: Sumatra: Sukaranda
1. Including Udamolcia Enderlein, 1920; the group is currently being revised by the junior author.
are almost entirely attributable to his giving an
inordinate amount of weight to the number of
metasomal tergites that have transverse sub-poste-
rior grooves. Only rarely does this character have
any significance at the genus level in the Braconi-
nae, but Enderlein based several of his new genera
solely on its distribution. It is perhaps not too sur-
prising therefore that only a few of his genera have
stood the test of time.
ANNOTATED CATALOGUE
The following is a list of all the species of Bra-
coninae described by Enderlein (1905a, 1905b,
1920) whose types are in the PAN collection.
Nearly all of these species were described in Ender-
lein's 1920 paper, however, the five species of
Rhamnura Enderlein were described in Enderlein
(1905a) and one species of Gastrotheca Guérin-
Méneville (not Fitzinger) (= Physaraia Shenefelt)
were described in Enderlein (1905b). The type
specimens of all of these are in the PAN except for
those of three of the species of Rhamnura and
paralectotypes of the remaining two, which are in
the Berlin Museum.
Enderlein’s original publications generally give
considerable detail concerning the type series and
the type localities. Where the type series includes
members of only one sex he usually placed a "Typus’
label on one individual and ‘Co-Typus’ labels on the
254
remainder thus giving a clear indication of which
specimen he wished the species concept to be based
upon; in nearly all such cases we have selected his
specimen bearing the ‘Type’ label as lectotype.
However, where the type series contains both
males and females he usually placed a ‘Type’ label
on one female and on one male; in these cases we
have generally selected the female as the lectotype
as this is the sex upon which most species descrip-
tions in the Braconidae are based. Because of Ender-
lein’s detailed notes on the type material, only a
summary is provided below; the type locality of the
primary types is given in table 1. It should be noted
that Shenefelt (1978) also gives some type locality
details corresponding to that of the holotype or
those specimens bearing ‘Type’ labels.
The species are listed below under their original
generic combinations in alphabetical order. Where
appropriate, notes are provided on the conditions
of the specimens, homogeneity of the type series
and previous taxonomic changes. The correct ge-
neric placement of most of the species studied is
given after the heading ‘Identity’; however, for
some it has not been possible to make a definitive
statement about their generic placement and in
these cases a few notes are usually given indicating
possible affinities. Their placement will be dis-
cussed in a forthcoming paper on the Indo-Austral-
ian genera of Braconinae by the junior author.
Antidiolcus excisus
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 female paralecto-
type not found, but should be present according to
the original description.
Notes. - This is the type species of Antidiolcus
Enderlein which was synonymized with Nesaulax
Roman by Quicke (1984a).
Identity. - Nesaulax excisus: Quicke (1984a).
Antiolcia grata
Type material. - Female holotype.
Notes. - On Enderlein’s type label this is labelled
as ‘Antiolcus gratus.
Identity. - Curriea grata comb. nov.
Antiolcia mitelligera
Type material. - One female with ‘Type’ label
hereby designated as lectotype and so labelled; one
male paralectotype also with ‘Type’ label.
Notes. — This is the type species of Antiolcia
Enderlein which was synonymized with Odontosca-
pus Kriechbaumer by Quicke (1981). Because this
taxon is aberrant in Odontoscapus it seems best to
retain it in Antiolcia till its position will be dis-
cussed in the forthcoming paper by the junior au-
thor. Both specimens are rather badly eaten-out but
the male is more badly damaged than the female.
The specimens are labelled by Enderlein as
‘Antiolcus mitelliger’.
Identity. - Anziolcia mitelligera: Quicke (1981).
Antiolcus apicalis
Type material. - Female holotype.
Notes. - This is the type species of Antiolcus
Enderlein which was synonymized with Hemz-
bracon Szépligeti by Quicke (1989c).
Identity. - Hemibracon apicalis: Quicke (1989c).
Antiolcus subfractus
Type material. - Female holotype.
Identity. - Digonogastra subfracta comb. nov.
Aphrastobracon gratiosus
Type material. - Male holotype.
Notes. - Placed by Enderlein in the Aphrastobra-
coninae which is now considered to be a tribe of
Braconinae.
Identity. - Curriea gratiosa comb. nov.
Aphrastobracon guttifer
Type material. - Male holotype.
Notes. - See A. gratiosus (above).
Identity. - Curriea guttifer comb. nov.
Bathyaulax laeviventris
Type material. - Female holotype.
Identity. - Bathyaulax laeviventris.
QUICKE & VAN ACHTERBERG: Enderlein’s Braconinae
Bathyaulax latiangulata
Type material. - Female holotype.
Notes. - The generic identity of this species is
hereby confirmed but no opinion is being given
concerning the validity or otherwise of the specific
synonymy proposed by Granger (1949).
Identity. — Bathyaulax foveiventris (Roman)
(synonymized by Granger, 1949).
Bathyaulax nigriceps
Type material. - Male holotype.
Identity. - Bathyaulax nigriceps.
Bathyaulax rugiventris
Type material. - Male holotype.
Identity. - Bathyaulax rugiventris.
Bracon merinotoides
Type material. - Female holotype.
Notes. - This species was transferred to Euwro-
bracon Ashmead by Fahringer (1927) and subse-
quently synonymized with E. forticornis (Cameron)
by Quicke (1989b).
Identity. - Euwrobracon forticornis (Cameron).
Campyloneurus albicans
Type material. - One female with "Typus’ label
hereby designated as lectotype; 1 ‘male’ paralecto-
type with ‘Typus’ label; 2 female and 2 male para-
lectotypes with ‘Co-Typus’ labels.
Notes. - The specimen labelled by Enderlein as
a male and bearing a ‘Typus’ label is in fact a female.
Identity. - Belongs to the Pachybracon Cameron-
group.
Campyloneurus basalis Enderlein (not Szépligeti)
Type material. - One female with ‘Typus’ label
hereby designated as lectotype; 1 male paralecto-
type with ‘Typus’ label; 11 female paralectotypes
with ‘Co-Typus’ labels.
Identity. - Campyloneurus enderleini Fahringer,
1931; replacement name.
Campyloneurus cincticauda
Type material. - Female holotype.
Identity. — Plesiobracon cincticauda comb. nov.
Campyloneurus cingulicauda
Type material. - Female holotype.
Identity. - Campyloneurus cingulicauda.
Campyloneurus disciventris
Type material. - One female with ‘Typus label
hereby designated as lectotype; 1 male paralecto-
type with ‘Typus’ label; 2 female and 6 male para-
lectotypes with ‘Co-Typus’ labels.
Identity. — Chelonogastra disciventris comb.
nov.
255
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Campyloneurus gibbiventris
Type material. - Female holotype.
Identity. - Belongs to the Pachybracon Cameron-
group.
Campyloneurus impressimargo
Type material. - One female with "Typus’ label
hereby designated as lectotype; 1 female paralecto-
type ‘Co-Typus' label.
Identity. - Campyloneurus impressimargo.
Campyloneurus latispeculum
Type material. - Female holotype.
Identity. - Campyloneurus latispeculum.
Campyloneurus maculistigma
Type material. - One female with “Typus’ label
hereby designated as lectotype; 1 female paralecto-
type ‘Co-Typus’ label, and belonging to Chaoilta.
Identity. - Uncertain.
Campyloneurus marginiventris
Type material. - Female holotype.
Identity. - Campyloneurus marginiventris.
Campyloneurus nigricosta
Type material. - One female with ‘Co-Typus’
label hereby designated as lectotype; 3 female para-
lectotypes with ‘Co-Typus’ labels and conspecific,
agreeing well with the original description. One
female has the basal half of the pterostigma yellow
and is not conspecific. Because this character was
not indicated in the original description this spec-
imen cannot be the lectotype, despite its ‘Typus’
label.
Identity. - Campyloneurus nigricosta.
Campyloneurus punctativentris
Type material. - Male holotype.
Identity. — Campyloneurus punctativentris; a
provisional identification because it is a male.
Campyloneurus reticulatus
Type material. - Female holotype.
Identity. - Campyloneurus reticulatus
Campyloneurus serenans
Type material. - Female holotype.
Notes. - The location of the type is not indicated
by Shenefelt (1978); it is in the PAN collection.
Identity. - Campyloneurus serenans.
Campyloneurus speculiger
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 4 female paralecto-
types with ‘Co-Typus’ labels.
Identity. - Campyloneurus speculiger.
256
Campyloneurus tibialis
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 2 female paralecto-
types with ‘Co-Typus’ labels.
Identity. - Campyloneurus tibialis.
Campyloneurus trispeculatus
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 male paralecto-
type with ‘Type’ label; 1 female paralectotype with
‘Co-Typus’ label.
Identity. - Campyloneurus trispeculatus.
Campyloneurus undicuneus
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 male paralecto-
type with Type’ label; 14 female and 5 male para-
lectotypes with ‘Co-Typus’ labels.
Identity. - Uncertain.
Craspedolcus fraternus
Type material. - One female with ‘Co-Typus’
label hereby designated as lectotype; 1 female para-
lectotype with ‘Type’ label. The lectotype is not
conspecific with the paralectotype and is the only
specimen agreeing with the original description
( 1. Tergit poliert glatt‘). The paralectotype be-
longs to Craspedolcus trisulcatus.
Identity. - Craspedolcus fraternus.
Craspedolcus obscuriventris
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 male paralecto-
type with ‘Type’ label and one of the other males
from Sukaranda must be also a paralectotype. The
other 3 are obviously added later.
Identity. - Craspedolcus obscuriventris.
Craspedolcus quadrirugulosus
Type material. - Female holotype.
Identity. - Isomecus quadrirugulosus comb. nov.
Craspedolcus strigidorsum
Type material. - Female holotype.
Identity. - Uncertain.
Craspedolcus trisulcatus
Type material. - One female with ‘Type’ label is
missing the ovipositor sheath, both left wings and
the right fore wing. Therefore we select and hereby
designate the other specimen with 'Co-Typus’ label
as lectotype. One female paralectotype with ‘Type’
label. Enderlein indicated in his description that he
had a larger and a smaller specimen, so his state-
ment at the end of the description that he had only
one female is obviously false.
Identity. - Craspedolcus trisulcatus.
Cyanopterus chilensis
Type material. - Male holotype.
Notes. — The holotype has the head severely
damaged, making its generic placement less certain
at present.
Identity. - Digonogastra chilensis; provisional
identification.
Cyanopterus filiseta
Type material. - Female holotype.
Identity. - Megabracon filiseta comb. nov.
Cyanopterus solox
Type material. - Female holotype.
Notes. - This species was not mentioned in the
recent revision of Exzrobracon Ashmead by Quicke
(1989b).
Identity. — Euurobracon triplagiata (Cameron)
syn. nov.
Cyanopterus thodeanus
Type material. - Female holotype.
Identity. - Compsobracon thodeanus comb. nov.
Diamblomera acuticella
Type material. - Female holotype.
Notes. - This is the type species of Diamblomera
Enderlein which was synonymised with Odontosca-
pus Kriechbaumer by Quicke (1981); however, we
now consider the two genera not to be synonymous
and therefore we treat Diamblomera provisionally
as a valid genus.
Identity. - Diamblomera acuticella.
Diamblomera grisescens
Type material. - Female holotype.
Identity. - Diamblomera grisescens.
Diamblomerina speciosa
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 2 female paralecto-
types with ‘Co-Typus’ labels.
Notes. - This is the type species of Diamblome-
rina Enderlein which is treated here as a new junior
synonym of Nedinoschiza Cameron.
Identity. - Nedinoschiza speciosa comb. nov.
Diolcia bicarinata
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 4 female paralecto-
types with ‘Co-Typus’ labels.
Notes. - This is the type species of Diolcıa En-
derlein which is treated here as a new junior syn-
onym of Campyloneurus Szépligeti.
Identity. — Campyloneurus bicarinatus comb.
nov.
QUICKE & VAN ACHTERBERG: Enderlein’s Braconinae
Diolcia limbaticauda
Type material. - Female holotype.
Identity. - Campyloneurus limbaticauda comb.
nov.
Ectemnoplax peruliventris
Type material. - Female lectotype (hereby desig-
nated; previously selected and labelled by C. van
Achterberg); 1 male paralectotype and a long series
of paralectotypes labelled as ‘Co-Typus’ by Ender-
lein.
Identity. - Ectemnoplax peruliventris.
Ectemnoplax lurida
Type material. - Female holotype.
Identity. - Monocoila lurida comb. nov.
Euryacria flavipera
Type material. - Female holotype.
Notes. - This is the type species of Euryacria
Enderlein which was synonymized with Bathyaulax
Szépligeti by Quicke (1981).
Identity. - Bathyaulax flavipera: Quicke (1981).
Gastrotheca sumatrana
Type material. - One female with ‘Type’ label
hereby designated as lectotype; the female paralec-
totype is missing.
Notes. - Only the lectotype was examined by
Donaldson (1989).
Identity. - Physarata sumatrana:
(1978).
Shenefelt
Goniobracon denticornis
Type material. - Holotype female.
Identity. - Serraulax denticornis comb. nov.
Goniobracon gutta
Type material. - Female lectotype (designated by
Shenefelt 1978: 1684); 1 male paralectotype.
Notes. - Goniobracon Szepligeti is a junior syn-
onym of Bathyaulax Szépligeti (see Quicke 1981);
however, so many mistakes in the interpretation of
these genera have occurred in the past that it is
considered best to reclassify all species separately.
Identity. - Bathyaulax gutta comb. nov.
Goniobracon helvimacula
Type material. - Male holotype.
Identity. — Zaglyptogastra helvimacula comb.
nov.
Goniobracon pectinatus
Type material. - Female holotype.
Identity. - Bathyaulax pectinatus comb. nov.
257
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Hemibracon elegantulus
Type material. - One female from Sumatra with
Typus’ label hereby designated as lectotype; 10
male and 6 female paralectotypes are in PAN and
additional paralectotypes from Taiwan are in the
Deutsche Entomologisches Institut at Eberswalde.
Notes. - All the specimens in the type series
appear to be conspecific with one another. The
generic placement of this species is hereby con-
firmed but no statement is being made concerning
its synonymy with Stenobracon trifasciatus Szé-
pligeti, 1908 as proposed by Watanabe (1932) and
through that, with S. niceville: (Bingham, 1901).
Identity. - Stenobracon nicevillei (Bingham).
Hemibracon w-impressum
Type material. - Female holotype.
Identity. - Hemibracon w-impressus.
Iphiaulax sauteri
Type material. - Female lectotype with ‘Type!
label hereby designated; 1 female and 4 males pa-
ralectotypes with only lectotype label.
Identity. - Iphiaulax sauteri.
Iphiaulax udei
Type material. - Female holotype.
Identity. - [phiaulax udei.
Ipobracon acuticampus
Type material. - Female holotype; the head, the
major part of the mesosoma and part of the met-
asoma (including the ovipositor) missing because
of damage by dermestids.
Identity. - Digonogastra acuticampa comb. nov.
Ipobracon alboniger
Type material. - Female holotype.
Identity. - Digonogastra alboniger comb. nov.
Ipobracon angustisulca
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 15 female paralec-
totypes with ‘Co-Typus’ labels.
Identity. - Uncertain, close to I. laevibasis Ender-
lein (q.v.).
Ipobracon aureopilosus
Type material. - Female holotype.
Identity. - Digonogastra aureopilosa comb. nov.
Ipobracon basispeculum
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 female paralecto-
type with ‘Co-Typus’ label.
Notes. - Probably conspecific with I. sexrugosus
Enderlein (see below).
Identity. - Uncertain.
258
Ipobracon biareatus
Type material. - One female from Peru with
‘Type’ label hereby designated as lectotype; 1 female
paralectotype with ‘Co-Typus’ label is not conspe-
cific.
Identity. - Digonogastra biareata comb. nov.
Ipobracon bicristatus
Type material. - Female holotype.
Identity. - Digonogastra bicristata comb. nov.
Ipobracon bicuneatus
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 female paralecto-
type without ‘Co-Typus’ label.
Identity. - Digonogastra bicuneata comb. nov.
Ipobracon bicuneus
Type material. - Female holotype.
Identity. - Digonogastra bicunea comb. nov.
Ipobracon brevicapulus
Type material. - Female holotype.
Identity. - Digonogastra brevicapula comb. nov.
Ipobracon brevicuneus
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 3 female paralecto-
types with ‘Co-Typus’ labels and a further three
females without ‘Co-Typus’ labels but clearly in-
cluded in the type series by Enderlein (1920).
Identity. - Digonogastra brevicunea comb. nov.
Ipobracon cameroni
Type material. - Female holotype.
Identity. - Digonogastra cameroni comb. nov.
Ipobracon columbianus
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 female paralecto-
type with ‘Co-Typus’ label.
Identity. - Digonogastra columbiana comb. nov.
Ipobracon crassecrenulatus
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 female paralecto-
type with ‘Co-Typus’ label.
Identity. - Hemibracon crassecrenulatus comb.
nov.
Ipobracon crenulatus Enderlein (not Szépligeti)
Type material. - Female holotype.
Notes. - Given the replacement name crenulato-
rius by Shenefelt (1978: 1814).
Identity. - Digonogastra crenulatoria (Shenefelt)
comb. nov.
Ipobracon crista
Type material. - One female with "Typus’ label
hereby designated as lectotype; 1 male paralecto-
type with ‘Typus’ label; 38 female and 14 male
paralectotypes with 'Co-Typus’ labels.
Identity. - Hybogaster crista comb. nov.
Ipobracon digitatus
Type material. - Female holotype.
Identity. - Digonogastra digitata comb. nov.
Ipobracon duploareatus
Type material. - Female holotype.
Identity. - Digonogastra duploareata comb. nov.
Ipobracon ecuadorensis
Type material. - Female holotype.
Identity. - Digonogastra ecuadorensis comb. nov.
Ipobracon fenestratus
Type material. - Female holotype.
Identity. - Digonogastra fenestrata comb. nov.
Ipobracon flavicaligatus
Type material. - Female holotype.
Identity. - Digonogastra flavicaligata comb. nov.
Ipobracon laevibasis
Type material. - Female holotype.
Notes. - Appears to be closest to Digonogastra
Ashmead but may warrant placement in a separate
genus when that group is revised.
Identity. - Uncertain.
Ipobracon laeviventris
Type material. - Female holotype.
Notes. - The head is missing and therefore we
cannot be certain about the generic placement of
this species at present.
Identity. - Uncertain.
Ipobracon latecrenulatus
Type material. - Female holotype.
Identity. — Digonogastra latecrenulata comb.
nov.
Ipobracon latefasciatus
Type material. - Female holotype.
Identity. - Digonogastra latefasciata comb. nov.
Ipobracon laticampus
Type material. - Female holotype.
Identity. - Digonogastra laticampa comb. nov.
Ipobracon laticuneus
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 female paralecto-
type with ‘Co-Typus’ label.
Identity. - Digonogastra laticunea comb. nov.
QUICKE & VAN ACHTERBERG: Enderlein's Braconinae
Ipobracon latisulca
Type material. - One female with “Typus’ label
hereby designated as lectotype; 1 male paralecto-
type with “Typus label; 49 female and 3 male pa-
ralectotypes with ‘Co-Typus’ labels.
Identity. - Uncertain, probably close to Craspe-
dolcus Enderlein.
Ipobracon longicapulus
Type material. - Female holotype.
Identity. - Digonogastra longicapula comb. nov.
Ipobracon macellus
Type material. - Male holotype.
Identity. - Digonogastra macella comb. nov.
Ipobracon maculicosta
Type material. - One female from Java with
‘Type’ label hereby designated as lectotype and has
been so labelled; 1 male paralectotype also with
‘Type’ label; 5 female and 9 male paralectotypes
with ‘Co-Typus’ labels.
Notes. - The male with the ‘Type’ label may not
be conspecific with the lectotype female because it
has forewing vein cu-a more or less interstitial,
hindwing vein 1r-m shorter than vein SC+R1 and
a posteriorly simple propodeum. The lectotype be-
longs to the Craspedolcus group having the poste-
rior margin of the propodeum ‘pinched-up’ in the
middle and forewing vein cu-a distinctly postfurcal
(see Quicke 1984b).
Identity. - Craspedolcus maculicosta comb. nov.
Ipobracon magnificus
Type material. - Female holotype.
Notes. - Transferred to Lasiophorus by Fahrin-
ger (1931); however, Plaxopsts, the genus to which
it belongs, is distinct from Lasiophorus despite the
fact that both have a strong facial protuberance (van
Achterberg 1983).
Identity. - Plaxopsis magnificus comb. nov.
Ipobracon mediofuscus
Type material. - Female holotype.
Identity. - Digonogastra mediofusca comb. nov.
Ipobracon nigripalpalis
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 female paralecto-
type with ‘Co-Typus’ label.
Notes. — This species was transferred to Atany-
colus Foerster by Shenefelt (1978); Hemibracon
Szépligeti is closely related to Atanycolus.
Identity. - Hemibracon nigripalpalis comb. nov.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Ipobracon nigripectus
Type material. - One female with ‘Type’ label
hereby designated as lectotype; one paralectotype
female with ‘Co-Typus’ label.
Identity. - Digonogastra nigripecta comb. nov.
Ipobracon obtusicampus
Type material. - Female holotype.
Notes. - The head, propleuron and ovipositor are
missing.
Identity. — Digonogastra obtusicampa comb.
nov.
Ipobracon ochripes
Type material. - Female holotype.
Identity. - Digonogastra ochripes comb. nov.
Ipobracon parvispeculum
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 2 paralectotype fe-
males with ‘Co-Typus’ labels.
Identity. - Uncertain, appears to be closely re-
lated to I. sexrugosus (q.v.).
Ipobracon penniseta
Type material. - Female holotype.
Identity. - Digonogastra penniseta comb. nov.
Ipobracon rectivena
Type material. - Female holotype.
Identity. - Digonogastra rectivena comb. nov.
Ipobracon rugiventris
Type material. - One female with ‘Type’ label
hereby designated as lectotype; one paralectotype
female with ‘Co-Typus’ label.
Identity. — Spathulibracon rugiventris comb.
nov.
Ipobracon sexrugosus
Type material. - Male holotype.
Identity. - Uncertain.
Ipobracon sigillatus
Type material. - Female holotype.
Identity. - Digonogastra sigillata comb. nov.
Ipobracon speculatus
Type material. - Female holotype.
Identity. - Digonogastra speculata comb. nov.
Ipobracon trirugosus
Type material. - Female holotype.
Notes. - The location of the type specimen is not
indicated in Shenefelt (1978); it is in the PAN
collection.
Identity. - Uncertain.
Ipobracon tuberculatus
Type material. - Female holotype.
Identity. - Digonogastra tuberculata comb. nov.
Megagonia breviseta
Type material. - One female with "Typus’ label
hereby designated as lectotype; 1 male paralecto-
type with "Typus’ label; 1 female paralectotype with
‘Co-Typus’ label.
Identity. - Iphiaulax breviseta comb. nov.
Megagonia griseiseta
Type material. - Female lectotype hereby desig-
nated; one female paralectotype.
Identity. - Zaglyptogastra griseiseta comb. nov.
Megagonia plumiseta
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 paralectotype fe-
male with ‘Co-Typus’ labels.
Notes. - The lectotype is in poor condition with
no head or anterior metasoma. The other specimen
with ‘Co-Typus’ label was not selected, since, al-
though its head is present, the mesosoma is dam-
aged.
Identity. - Zaglyptogastra plumiseta comb. nov.
Merinotus bispeculum
Type material. - One female with ‘Type’ label
hereby designated as lectotype; four paralectotype
males, one with ‘Type’ and three with ‘Co-Typus’
labels.
Identity. - Shelfordia bispeculum comb. nov.
Merinotus criniseta
Type material. - One female with ‘Typus’ label
hereby selected as lectotype and so labelled; one
male paralectotype with ‘Type’ label; 3 female pa-
ralectotypes with ‘Co-Typus’ labels.
Identity. - Shelfordia criniseta comb. nov.
Merinotus flagriseta
Type material. - One female with ‘Type’ label
hereby designated as lectotype; sixteen paralecto-
type females with ‘Co-Typus’ labels.
Additional material. — Five males det. as fla-
griseta by Enderlein but without ‘Co-Typus’ labels.
Notes. - No opinion is being given here as to
whether the males that were tentatively identified
by Enderlein are conspecific with the females of the
type series.
Identity. - Shelfordia flagriseta comb. nov.
Merinotus gibber
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 106 paralectotype
females with 'Co-Typus’ labels.
Notes. — All the specimens in the type series
appear to be conspecific; the type series is currently
split between two cabinet drawers.
Identity. - Cratobracon gibber comb. nov.
Merinotus gracilis
Type material. - Holotype.
Notes. - The metasoma is missing but Ender-
lein’s label and original description indicate that it
is a female.
Identity. - Nesaulax gracilis comb. nov.
Merinotus ingentiseta
Type material. - One female with ‘Type’ label
hereby selected as lectotype; one female paralecto-
type with 'Co-Typus’ label.
Identity. - Shelfordia ingentiseta comb. nov.
Merinotus laticampus
Type material. - One female with ‘Type’ label
hereby selected as lectotype and so labelled; one
male paralectotype with ‘Type’ label; fourteen fe-
male paralectotypes with ‘Co-Typus’ labels.
Notes. - The lectotype and the paralectotype
male are not congeneric; the paralectotype male
belongs to the genus Shelfordia Cameron.
Identity. - Gronaulax laticampus comb. nov.
Merinotus nigriseta
Type material. - One female with ‘Type’ label
hereby selected as lectotype and so labelled; eight
female paralectotypes with ‘Co-Typus’ labels.
Identity. - Gammabracon nigriseta comb. nov.
Merinotus quadricarinatus
Type material. - One female with ‘Type’ label
hereby selected as lectotype; 1 ‘male’ paralectotype
with a ‘Type’ label; 89 female paralectotypes with
‘Co-Typus’ labels.
Notes. - The ‘male’ paralectotype is in fact a
female with the ovipositor broken off near its base.
Identity. - Shelfordia quadricarinata comb. nov.
Merinotus rimicuneus
Type material. - One female with ‘Type’ label
hereby selected as lectotype and so labelled; one
male paralectotype with ‘Type’ label; 2 female and
1 male paralectotypes with ‘Co-Typus’ labels.
Identity. - Shelfordia rimicunea comb. nov.
Merinotus ruficauda
Type material. - Female holotype.
Identity. - Rhadinobracon ruficauda comb. nov.
QUICKE & VAN ACHTERBERG: Enderlein’s Braconinae
Merinotus trispeculum
Type material. - One female with ‘Type’ label
hereby designated as lectotype; four paralectotype
males, one with ‘Type’ and three with ‘Co-Typus’
labels.
Identity. - Uncertain.
Monolcia angulosa
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 5 female paralecto-
types with ‘Co-Typus’ labels.
Identity. - Campyloneurus angulosus comb. nov.
Monolcia apicalis
Type material. - One female with ‘Type’ label
hereby designated as lectotype and so labelled; one
male paralectotype with ‘Type’ label; three female
paralectotypes with ‘Co-Typus’ labels.
Identity. — Belongs to Pachybracon Cameron-
group.
Monolcia consimilis
Type material. - 1 female with ‘Type’ label here-
by designated as lectotype; 1 female paralectotype
with 'Co-Typus’ label.
Identity. - Uncertain.
Monolcia flavicosta
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 paralectotype fe-
male with ‘Co-Typus’ label.
Notes. - The lectotype has the metasoma glued
on to the mesosoma.
Identity. — Provisionally included in Campylo-
neurus despite its apically truncate scapus. Campy-
loneurus flavicosta comb. nov.
Monolcia flavimarginata
Type material. - Female holotype.
Identity. - Uncertain.
Monolcia serenimanus
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 paralectotype fe-
male with ‘Co-Typus’ label.
Notes. - The lectotype and paralectotype do not
appear to be conspecific, the paralectotype having
a preapical dorsal nodus and apico-ventral serra-
tions on the ovipositor, both of which are absent in
the lectotype.
Identity. - Uncertain.
Monolcia tricarinata
Type material. - Female holotype.
Notes. — This is the type species of Monolcia
Enderlein which is hereby synonymized with Cam-
pyloneurus Szépligeti.
Identity. — Campyloneurus tricarinatus comb.
nov.
261
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Odontopygia tridentata
Type material. - One female with "Typus’ label
hereby designated as lectotype; 2 female paralecto-
types with ‘Co-Typus’ labels.
Notes. - This is the type species of Odontopygia
Enderlein which was synonymized with Trigas-
trotheca Cameron by Quicke (1987).
Identity. — Trigastrotheca tridentata: Quicke
(1987).
Plagiozina pravivena
Type material. - One female with ‘Type’ label
hereby designated as lectotype; one male paralecto-
type with ‘Type’ label; eight female and one male
paralectotypes with ‘Co-Typus’ labels.
Notes. - This is the type species of Plagiozina
Enderlein which was synonymized with Nesazlax
Roman by Quicke (1984a) and is most likely a
synonym of N. flagellaris Roman, 1913.
Identity. - Nesaulax pravivena: Quicke (1984a)
Platybracon carinicornis
Type material. - One female with ‘Typus’ label
hereby designated as lectotype; 5 female paralecto-
types with ‘Co-Typus’ labels.
Notes. - Platybracon Szépligeti is a junior syn-
onym of Chaoilta Cameron (see Roman 1913,
Quicke 1987); however, the two were dealt with
separately in Shenefelt's (1978) catalogue.
Identity. - Chaoılta carinicornis comb. nov.
Platybracon celebensis
Type material. - Female holotype.
Notes. - See P. carinicornis (above). Belongs to
the decorata group of species having a horn-like
projection above the plate-like facial projection.
Identity. - Blastomorpha celebensis comb. nov.
Platybracon comptus
Type material. - One female with Typus’ label
hereby designated as lectotype; 1 female paralecto-
type with 'Co-Typus’ label.
Notes. - See P. carinicornis (above).
Identity. - Chaoılta compta comb. nov.
Pseudospinaria sannio
Type material. - Female holotype.
Notes. - Erroneously included in the Exothecinae
by Enderlein (1920: 54); Pseudospinaria Enderlein
is also a member of the Braconinae.
Identity. — Trispinaria sannio comb. nov.
Rhadinobracon rhyssides
Type material. - Female holotype.
Identity. - Ischnobracon rhyssides comb. nov.
Rhamnura capillicauda
Type material. - Lectotype female (designated by
Shenefelt 1978: 1719); 4 female and 1 male para-
lectotype are in the Museum fiir Naturkunde in
Berlin.
Notes. - Redescribed and illustrated by van Ach-
terberg (1981).
Identity. - Rhamnura capillicauda.
Rhamnura filicauda
Type material. - Lectotype female (designated by
Shenefelt 1978: 1719); 2 paralectotype females are
in the Museum fiir Naturkunde in Berlin.
Identity. - Rhamnura filicauda.
Syntomernus pusillus
Type material. - One female with ‘Type’ label
hereby designated as lectotype and so labelled; one
‘male’ paralectotype with ‘Type’ label; 5 female pa-
ralectotypes with ‘Co-Typus’ labels.
Notes. - The ‘male’ paralectotype has no meta-
soma and therefore its sex cannot be verified.
Identity. - Syntomernus pusillus.
Udamolcia albescens
Type material. - One female with “Typus’ label is
damaged, therefore we designate hereby one of the
two females with ‘Co-Typus’ label as lectotype; 2
female paralectotypes one with ‘Type’and one with
‘Co-Typus’ label.
Notes. - This is the type species of Udamolcia
Enderlein which was synonymized with Pachy-
bracon Cameron by Quicke (1984c).
Identity. - Belongs to the Pachybracon-group.
Udamolcia albimanus
Type material. - Male holotype. In the collection
under Campyloneurus.
Identity. - Uncertain.
Udamolcia denticornis
Type material. - One female with “Typus label
hereby designated as lectotype; 22 female paralec-
totypes with ‘Co-Typus’ labels.
Identity. - Uncertain; a member of the Atanyco-
lus Foerster group (see Quicke 1987).
Udamolcia gracilis
Type material. - Male holotype. In the collection
under Campyloneurus.
Identity. - Uncertain. Belongs to the Campylo-
neurus-group.
Udamolcia insolita
Type material. - Female holotype. In the collec-
tion under Campyloneurus.
Identity. — Belongs to the Pachybracon-group.
Udamolcia jucunda
Type material. - Female holotype.
Identity. - Undabracon jucundus comb. nov.
Udamolcia laqueata
Type material. - One female with ‘Typus’ label
hereby designated as lectotype; 1 male paralecto-
type with “Typus’ label; 3 female and 1 male para-
lectotypes with ‘Co-Typus’ labels.
Identity. - Cordibracon laqueatus comb. nov.
Udamolx aterimma
Type material. - Female holotype.
Identity. - Digonogastra aterimma comb. nov.
Udamolx atricauda
Type material. - Male holotype.
Identity. - Archibracon atricauda: Quicke (1989a).
Udamolx catharinensis
Type material. - One female with ‘Type’ label
hereby designated as lectotype; 1 female and 1 male
paralectotypes with ‘Co-Typus’ labels and 1 male
paralectotype without 'Co-Typus’ label.
Identity. — Hemibracon catharinensis comb.
nov.
Udamolx gutta
Type material. - Female holotype.
Notes. - This is the type species of Udamolx
Enderlein which was synonymized with Archi-
bracon Saussure by Quicke (1983).
Identity. — Archibracon cameroni Brues (syn-
onymized by Quicke 1989a).
Udamolx polita
Type material. - Female holotype.
Identity. - Hemibracon politus comb. nov.
Udamolx vaucristatum
Type material. - 1 female with ‘Type’ label is the
holotype. The male (with only Enderlein’s identi-
fication label) was excluded from the type series by
Enderlein (1920: 100).
Identity. - Hemibracon vaucristatus comb. nov.
ACKNOWLEDGEMENTS
We would like to thank Professor Dr B. Pisarski,
Dr E. Kierych, Mr P. Marczak (PAN) and Magda
Kierych for their help and hospitality during our
visits to Warsaw and Lomna. Partly this research
was made possible by a Royal Society short term
exchange awarded to the senior author.
REFERENCES
Achterberg, C. van, 1981. The position of the genus
Rhamnura Enderlein among the Braconidae, Hyme-
QUICKE & VAN ACHTERBERG: Enderlein's Braconinae
noptera). — Entomologische Berichten, Amsterdam
41: 88-91.
Achterberg, C. van, 1983. Six new genera of Braconinae
from the Afrotropical Region (Hymenoptera, Braco-
nidae). — Tijdschrift voor Entomologie 126: 175-202.
Donaldson, J. S., 1988. Revision of the braconid genus
Physaraia (Hymenoptera, Ichneumonoidea). — Syste-
matic Entomology 14: 15- 39.
Enderlein, G., 1905a. Rhamnura, eine neue Braconiden-
gattung mit ausserordentlich langem Legrohr des
Weibchens. - Zoologischer Anzeiger 29: 195-198.
Enderlein, G., 1905b. Neue Braconiden aus dem indischen
und afrikanischen Gebiet. — Stettiner Entomologische
Zeitung 66: 227-236.
Enderlein, G., 1920. Zur Kenntnis aussereuropäischer
Braconiden. - Archiv für Naturgeschichte 84(A) (11):
51-224.
Fahringer, 1931. Opuscula braconologica. 2. Aethiopi-
schen Region. Lieferung 5: 305-384.
Granger, Ch., 1949. Braconides de Madagascar. — Mé-
moires de l'Institut Scientifique de Madagascar, A) 2:
1-428.
Quicke, D. L. J., 1981. A reclassification of some Oriental
and Ethiopean species of Braconinae (Hymenoptera:
Braconidae). - Oriental Insects: 14: 493-498.
Quicke, D. L. J., 1983. The Afrotropical genus Archibracon
Saussure (Hymenoptera: Braconidae: Braconinae):
Characteristics and new generic synonymy. - Entomol-
ogist’s monthly magazine 119: 147-150.
Quicke, D. L. J., 1984a. Two new genera of Afrotropical
Braconinae (Hymenoptera: Braconidae) with a partial
review of those genera with ‘merinotoid’ metasomas.
— Entomologist's Monthly Magazine 120: 37-45.
Quicke, D. L. J., 1984b. Redescription of Craspedolcus
with notes on included species and related genera
(Hymenoptera: Braconidae). — Oriental Insects 18:
354-360.
Quicke, D. L. J., 1984c. Further reclassification of Afro-
tropical and Indo-Australian Braconinae (Hymenop-
tera: Braconidae). - Oriental Insects 18: 339-353.
Quicke, D. L. J., 1987. The Old World genera of Braconine
wasps (Hymenoptera: Braconidae). — Journal of Nat-
ural History 21: 43-157.
Quicke, D. L. J., 1989a. Parasitic braconine wasps of the
genus Archibracon (Hymenoptera: Braconidae). —
Journal of Natural History 23: 29-70.
Quicke, D. L. J., 1989b. The Indo-Australian and E. Pa-
laearctic braconine genus Ezurobracon (Hymenop-
tera: Braconidae: Braconinae). — Journal of Natural
History 23: 775-802.
Quicke, D. L. J., 1989c. Reclassification of some New
World species of Braconinae (Hym., Braconidae). —
Entomologist's monthly magazine 125: 119-121.
Roman, A., 1913. Philippinische Schlupfwespen aus dem
schwedischen Reichsmuseum. I. - Arkiv för Zoologi 8:
7-59.
Shenefelt, R. D., 1978. Hymenopterorum Catalogus (nov.
ed.). Part 15. Braconidae, vol. 10. - The Hague, Junk,
pp 1425-1865.
Watanabe, C., 1932. A revision of Braconid species para-
sitic in the injurious insects of rice plant and sugar cane
in Japan and Formosa. — Transactions of the Sapporo
Natural History Society 12: 63-72.
Received: 14 September 1990
Revised version accepted: 25 October 1990
263
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
INDEX TO SPECIES GROUP NAMES
258
257
262
255
262
258
acuticampus
acuticella
albescens
albicans
albimanus
alboniger
angulosa
angustisulca
apicalis (Antiolcus)
apicalis (Monolcia)
aterrima
atricauda
aureopilosus
basalis
basispeculum
biareatus
bicarinata
bicristatus
bicuneatus
bicuneus
bispeculum
brevicapulus
brevicuneus
breviseta
cameroni
capillicauda
carinicornis
cariniventris
catharinensis
celebensis
chilensis
cincticauda
cingulicauda
columbianus
comptus
consimilis
crassecrenulatus
crenulatus
criniseta
crista
denticornis (Goniobracon)
denticornis (Udamolcia)
digitatus
disciventris
duploareatus
ecuadorensis
elegantulus
EXCISUS
fenestratus
filicauda
257
260
259
261
261
257
256
261
256
261
262
255
255
260
257
257
263
255
DI
256
261
262
263
259
255
259
263
259
259
255
259
261
259)
256
259
257
259
ZO]
259
259
256
259
256
259
255
255
255
256
259
260
filiseta
flagriseta
flavicaligatus
flavicosta
flavimarginata
flavipera
fraternus
gibber
gibbiventris
gracilis (Merinotus)
gracilis (Udamolcia)
grata
gratiosus
griseiseta
grisescens
gutta (Goniobracon)
gutta (Udamolx)
guttifer
helvimacula
impressimargo
ingentiseta
insolita
jucunda
laevibasis
laeviventris (Bathyaulax)
laeviventris (Ipobracon)
laqueata
latecrenulatus
latefasciatus
latiangulata
laticampus (Ipobracon)
laticampus (Merinotus)
laticuneus
latispeculum
latisulca
limbaticauda
longicapulus
lurida
macellus
maculicosta
maculistigma
magnificus
marginiventris
mediofuscus
merinotoides
mitelligera
nigriceps
nigricosta
nigripalpalis
nigripectus
nigriseta
obscuriventris
obtusicampus
ochripes
parvispeculum
pectinatus
penniseta
peruliventris
plumiseta
polita
pravivena
punctativentris
pusillus
quadricarinatus
quadrirugulosus
rectivena
reticulatus
rimicuneus
rhyssides
ruficauda
rugiventris (Bathyaulax)
rugiventris (Ipobracon)
sannio
sauteri
serenans
serenimanus
sexrugosus
sigillatus
solox
speciosa
speculatus
speculiger
strigidorsum
subfractus
sumatrana
tibialis
thodeanus
tricarinata
tridentata
trirugosus
trispeculatus
trispeculum
trisulcatus
tuberculatus
udei
undicuneus
vaucristatum
w-impressum
R. T. A. SCHOUTEN
Museon, Den Haag
A NEW SPECIES OF EUCHROMIUS GUENEE, 1845
(LEPIDOPTERA: PYRALIDAE: CRAMBINAE)
FROM AUSTRALIA
Schouten, R. T. A., 1990. A new species of Ewchromius Guenée, 1845 (Lepidoptera: Pyra-
lidae: Crambinae) from Australia. - Tijdschrift voor Entomologie 133: 265-267, figs. 1-5.
[ISSN 0040-7496]. Published 14 December 1990.
Euchromius cornus sp. n. is described from Australia. The external characters, male and
female genitalia are described and figured in detail.
R. T. A. Schouten, Museon, Dept. of Biology, Stadhouderslaan 41, 2517 HV Den Haag, The
Netherlands.
Key words. - Pyralidae; Crambinae; Euchromius, Australia; new species.
Presumably in 1991, a checklist of the Lepido-
ptera of Australia will be published. The only spe-
cies of Ewchromius occurring in Australia is des-
cribed in this article to have the name available for
this checklist. A revision of the other species will be
published shortly.
RTAS refers to author's collection, other abbre-
viations (codens) used in the list of type material
follow Arnett & Samuelson (1986). The terminol-
ogy used in the descriptions follows Schouten
(1988).
Euchromius cornus sp. n.
(figs. 1-5)
Type material. - Holotype: 8, ‘Sherlock R. W. Australia.
E. Clements. 98-188’, genitalia slide 17616 BM (BMNH).
- Paratypes: 1 8, Alexandria (BMNH); 1 8, Comrav
(BMNH); 2 6, 4 9, Broken Hill (RTAS, SAMA); 1 8, to
Alhannau above Column (SAMA); 1 d, Stevenson River
(SAMA); 2 8, 1 9, 3 km. NW by W. of Millstream HS.
(ANIC); 1 4, 2 9, 1 km. NE of Millstream HS. (ANIC,
RTAS); 1 g, 1 9, 1 km. N of Millstream HS. (ANIC,
RTAS); 1 9, 1/2 km. WNW of Millstream HS. (ANIC);
2 6, 1km NNE of Millstream WA (ANIC, RTAS); 1 4,
Camooweal (ANIC); 1 9, 31 km NNW Longreach, Q.
(ANIC); 1 6, Kimberley Research Stn Via Wyndham N.
W. Aust. (ANIC); 2 6, Ivanhoe. W. A (ANIC, RTAS); 1
8, Moora W Q (ANIC); 1 9, Limestone Creek, 42 km
NNW of Boulia, Q. (ANIC); 1 9, JimJim Creek 19 km
WSW of Mt. Cahill, N. T. (ANIC); 1 @, Charleville Q.
(ANIC); 2 6,1 9, Injune Q. (ANIC, RTAS); 1 9, Cunnam-
ula Q. (ANIC); 1 @, 1 9, Adaville Q. (ANIC); 1 9, Mil-
merran Q. (ANIC).
Diagnosis. - Externally almost indistinguishable
from Euchromius species with a double medial fas-
cia and frons with one point. Differs in male geni-
talia from all species in having the uncus armed
with a short, double, dorsal projection. Differs in
female genitalia from E. geminus, E. californicalis,
E. limaellus, E. ocelleus, E. saltalis, E. mythus and
E. tanalis in having a short ductus bursae without
clear sclerotizations.
External characters (fig. 1). - Wingspan 16-22
mm. Frons produced forward, conical with clear
point, creamy white, no ventral ridge; vertex
creamy white; labial palp two to two and a half,
sides creamy white at base, becoming light brown,
creamy white from above and below; maxillary palp
creamy white to light brown, dark ringed at base of
last segment; antenna creamy white. Thorax
creamy white to light brown; patagia creamy white;
tegulae creamy white, evenly mottled. Forewing,
Figs. 1-2. Euchromius cornus. 1, Head, lateral aspect,
scales removed, scale 1 mm; 2, Male, sclerotization of
tergite VIII, scale 1 mm.
265
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Fig. 3. Euchromius cornus, male genitalia, holotype; ventro-caudal aspect, valvae spread; aedeagus separated, lateral
aspect, slide BM 17616, scale 0.5 mm.
groundcolour creamy white densely suffused with
ochreous brown to brown scales; medial fascia dou-
ble, arched to nearly straight, running to one-sixth
to one-fifth of the dorsum; subterminal line ochre-
ous brown, inconspicuous, slightly closer to termi-
nation of posterior area than to terminal dots,
sometimes about halfway between terminal dots
and termination of posterior area; area adjacent to
terminal dots white; eight or nine black terminal
dots, formula 2-2-3-1 or 1-2-3-2 or 2-2-3-2; fringes
shiny, creamy white with two to three brown lines.
Hindwing creamy white, subterminal fascia absent,
termen darkly bordered; fringes white with creamy
white line.
266
Tergite VIII (fig. 2). - Tergite normally sclerot-
ized, posterior part triangular, stalk only partly vis-
ible, anterior pattern not visible.
Male genitalia (fig. 3). - Uncus with a short,
double, dorsal projection; gnathos longer, with dor-
sal dagger-shaped projections at each side of base,
two dorsal thorns relatively large, terminal part
long; tegumen without appendix angularis; saccu-
lus normal, processus of sacculus absent, processus
basalis long strongly sclerotized, two processi infe-
riores valvae, cucullus club-shaped, bent upward;
juxta triangular, armed with two dorsal projections;
vinculum small; aedeagus normal sized, one dentate
cornutus anterior of anellus connection, second,
Fig. 4. Euchromius cornus, female genitalia; ventral as-
pect, slide BM 17469, scale 0.5 mm.
SCHOUTEN: New Euchromius
------I- AL
A ><
si \ ayy) ©
U
\
\
t
Fig. 5. Distribution of Euchromius cornus.
fainter group of cornuti more posterior.
Female genitalia (fig. 4). - Papillae anales nor-
mal; membrane of tergite VIII with cord-like scle-
rotization, edges of tergite VIII not connected; os-
tium lip-shaped; ductus bursae short, broad under
ostium; ductus seminalis narrow; bursa copulatrix
oblong, two long signa, about equal in size.
Ecology. - Unknown. Two generations, the first
starts in February and ends at the end of May, most
specimens were caught in April and May. The se-
cond generation flies from August to the end of
November, most specimens in September and Oc-
tober.
Distribution (fig. 5). - Restricted to Australia,
where it can be found in most provinces.
REFERENCES
Arnett, R. H. & G. A. Samuelson, 1986. The insect and
spider collections of the world. 220 pp. - EJ.
Brill/Flora & Fauna publications, Gainesville.
Haworth, A. H., 1811. Lepidoptera Britannica, 3: 377-511.
— London.
Schouten, R. T. A., 1988. Revision of the species of the
genus Euchromius Guenée, 1845 (Lepidoptera: Pyra-
lidae: Crambinae) occurring in the Afrotropical region.
— Zoologische Verhandelingen Leiden 244: 1-64.
Received: 2 October 1990.
Accepted: 11 October 1990.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
BOOK ANNOUNCEMENTS AND REVIEWS
Larsen, T.B., 1990. The butterflies of Egypt. - Apollo
Books, Lundbyvej 36, DK-5700 Svendborg, Denmark:
1-112, figs. 1-7, tables 1-4, plates 1-8. [ISBN 87 88757
14 5, price 240 Danish Crowns].
The current knowledge of the butterflies of
Egypt is summarized for the first time since 1948.
All 58 species are depicted in full colour, together
with some photo's of the most characteristic hab-
itats. For each species the distribution inside and
outside Egypt is discussed together with their tax-
onomical status, ecology and habitat preferences.
Separate chapters deal with the ecological zones
of Egypt and its characteristic species, the biogeo-
graphy, migrant species, economic importance and
species expected to be found in Egypt in the near
future. This book is recommended warmly to eve-
ryone working on Mediterranean butterflies or in-
tending to visit Egypt.
[R. T. A. Schouten]
Merzheevskaya, O. I., 1989. Larvae of owlet moths (Noc-
tuidae). Biology, morphology, and classification. —
Amerind Publishing Co, New Delhi (outside India
exclusively marketed by E. J. Brill, Leiden etc): i-xx +
1-419, figs. 1-97. ISBN 90-04-08804-0. Price 115.-
Dutch guilders (appr. US $57.50).
This book is a translation of ‘Gusenitsy Sovok
(Noctuidae), ikh Biologiya i Morfologiya (Oprede-
litel’)’, which was published in 1967. It provides
information on the biology and morphology of
owlet moth larvae and keys to their subfamilies,
genera, and species. Morphological characters, body
coloration, and dermal patterns are detailed for
each species. The larvae of 144 species of owlet
moths are described and data on developmental
changes in instars given for 91 of them.
This book should be helpful to entomologists,
agricultural entomologists, plant-protection spe-
cialists, and teachers and students of biology, agri-
culture, and forestry.
[Information from the publisher]
Oman, P. W., W.J. Knight & M. W. Nielson, 1990. Leaf-
hoppers (Cicadellidae): A bibliography, generic check-
list and index to the world literature 1956-1985. - CAB
International Institute of Entomology, Wallingford: 1-
368. ISBN 0 85198 690 0. Price not given.
The book consists of a bibliography of more than
7000 titles for the period 1956-1985, an index to the
bibliography to guide users to specified subject-
matter fields and a complete check-list of generic
and family-group names used in the Cicadellidae.
Other important features are a review of cicadellid
literature for the period 1942-1955, a compilation
showing usage of family-group names in the taxo-
nomic literature during the years 1956-1985 and an
alphabetized list and grouping of subfamily and
tribal names within the family. The present bibli-
ography supplements Metcalf’s Bibliography of the
Cicadelloidea.
[From the introduction of this book].
Williams, D. J. & G. W. Watson, 1990. The scale insects
of the Tropical South Pacific region. Part 3. The Soft
Scales (Coccidae) and other families. - CAB Interna-
tional Institute of Entomology, Wallingford, UK: 1-
267, figs. 1-71. ISBN 0 85198 659 5. Price not given.
A continuation of the series started by the same
authors in 1988 (cf. Tijdschrift voor Entomologie
132: 114). The families discussed in the present
volume are Margarodidae, Ortheziidae, Eriococci-
dae, Dactylopiidae, Coccidae, Cerococcidae, Lecano-
diaspididae, Kerriidae, Asterolecaniidae, Conchas-
pididae and Halimococcidae. Keys are provided to
the families and genera, and many new species are
described and illustrated.
[From the abstract |.
JAN H. STOCK
Institute of Taxonomic Zoology, University of Amsterdam
A NEW SPECIES OF AUSTRODECUS
(PYCNOGONIDA) FROM NEW SOUTH WALES,
AUSTRALIA
Stock, Jan H., 1990. A new species of Austrodecus (Pycnogonida) from New South Wales,
Australia. — Tijdschrift voor Entomologie 133: 269-272, figs. 1-9. [ISSN 0040-7496]. Pub-
lished 14 December 1990.
Austrodecus staplesi sp.n. is the first member of the genus described from continental
Australia (New South Wales). The genus is mainly Gondwanian in its distribution, but for
three species recorded from the western belt of the Pacific Ocean.
Dr. J. H. Stock, Institute of Taxonomic Zoology, University of Amsterdam, P.O. Box 4766,
1009 AT Amsterdam, The Netherlands.
Key words. - Austrodecus, Pycnogonida; New Species; New South Wales; biogeography.
The genus Awstrodecus s.str. counts actually 24
named and 5 unnamed species (Stock 1957, 1968,
Hedgpeth & McCain 1971, Clark 1972a, b, Child
1983, 1988, Pushkin 1977). The genus was briefly
mentioned, without specific identification, in a pop-
ular paper by Staples (1977: 123) from ‘southern
Australian waters ... [where it] may be associated
with the hydroid Halicorniopsis elegans’. With the
exception of one dubious record of A. glaciale
Hogdson, 1907, off Tasmania (see Gordon 1944: 6,
Stock 1957: 46), there are no other Australian re-
cords, and as far as I know Staples’ Australian mate-
rial never has been formally described.
Through the courtesy of Dr. Harry A. ten Hove,
Amsterdam, I received a single specimen, fortu-
nately a male, of an Austrodecus collected on the
coasts of New South Wales. It represents a new
species, described in this paper.
Family AUSTRODECIDAE Stock, 1954
Genus Austrodecus Hodgson, 1907
Austrodecus staplesi sp.n.
(figs. 1-8)
Type material. - 1 8 (holotype), Australia, New South
Wales, Split Solitary Island, near Coff's Harbour; depth
12-14 m; rocky area with small caves, some corals, algae
and some sand; 26 Apr. 1986; leg. H. A. ten Hove, P.
Hutchings & R. Phipps (Zoölogisch Museum Amsterdam
Ba. 3322):
Description
Holotype 8. - The new species belongs to the
gordonae-section (Stock 1957) of the genus, char-
acterized by a 4-segmented oviger and the absence
of auxiliary claws.
Trunk (figs. 1-2) with 4 tall mid-dorsal spurs, one
on each trunk segment. Ocular tubercle taller than
trunk spurs, but rather plump; eyes well-pig-
mented. Less tall dorsal spurs on coxa 1 of legs 1 and
4 (1 spur), coxa 1 of legs 2 and 3 (2 spurs), and coxa
3 of legs 1 to 4 (1 spur). Abdomen (fig. 1) tuber-
culate, slightly overreaching coxa 3 of leg 4.
Palp (fig. 3) 5-segmented, segment 5 indistinctly
subdivided, bifid (fig. 4). Oviger 4-segmented, 1
spine on segment 3, 2 median and 4 distal spines on
segment 4 (fig. 5).
Legs (fig. 6) with a long dorsodistal seta on femur
and tibiae 1 and 2. Femur longest segment. Femoral
cement gland ventral, situated slightly before mid-
dle of segment; discharging through long duct (fig.
7). Other glands open through slightly raised pores
(fig. 8) on dorsal surface of tibia 1 (4 pores) and tibia
2 (2 pores). Propodus strongly curved; sole with c.
6 spinules; no auxiliary claws. As far as I can ascer-
tain, male genital pores situated on ventral surface
of coxa 3 (!) of all (!) legs.
Measurements of holotype (in um). - Length
trunk (tip ocular tubercle to tip abdomen) 1556:
length cephalic segment 653; width across 2nd lat-
eral processes 539; length abdomen 406; length
proboscis 1099.
Third leg: first coxa 144; second coxa 144; third
coxa 101; femur 376; first tibia 264; second tibia
241; tarsus 49; propodus 269; claw 120.
Remarks
Within the gordonae-section, the new species
269
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Figs. 1-8. Austrodecus staplesi n.sp., & holotype. 1, body, dorsal; 2, body, from the left; 3, palp; 4, distal part of palp;
5, oviger; 6, third leg; 7, femoral cement gland; 8, gland apertures on dorsal surface of tibia 2 of leg 3.
differs from A. frigorifugum Stock, 1954, A. stocki
Child, 1988, and A. oblongum Pushkin, 1977 (ge-
neric status of the latter uncertain because of aber-
rant shape of proboscis) in the absence of dorso-
distal femoral spurs. A. gordonae Stock, 1954, has
low mid-dorsal trunk tubercles, an almost straight
and shorter propodus, and lacks a femoral cement
gland tube. A. palazense Child, 1983 has a longer
distal tubercle on palp segment 2, as well as tuber-
culate and widely separated lateral trunk processes;
the oviger of this species is unknown.
As point of fact, only three species of
Austrodecus share with the new species the pres-
ence of a tubiform cement gland aperture located in
the middle part of the ventral surface of femur. Two
of these, A. aconae (Hedgpeth & McCain, 1971),
originally described as a species of Pantopipetta,
270
and A. kelpi Pushin, 1977 differ from the new spe-
cies in the absence or poor development of mid-
dorsal trunk spurs, and in the axial implantation of
the distal palp segment. The cther species,
A. tubiferum Stock, 1957, shows no doubt the
greatest resemblance to the new species.
A. tubiferum is known from Sagami Bay (Stock
1954, 1957) and Okinawa (Child 1988), and has a
similar pattern of spurs on trunk and legs. The new
species differs from A. tubiferum in having longer
mid-dorsal trunk spurs, a more strongly curved pro-
podus, a slightly different configuration of the distal
palp segments, and less slender legs (especially on
the level of tibia 2). The propodus of A. tubiferum
is shorter than tibia 2, that of A. staples: longer
than tibia 2.
The new species differs from all other members
NS
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STOCK: A new species of Austrodecus
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Fig. 9. Global reconstruction at 100 Ma B.P., after Smith & Briden, 1977. Dots indicate the actual distribution of the
24 named and 5 unnamed species of the genus Austrodecus (some species known from more than one locality, some
localities with more than one species). With the exception of the Japanese, Palau and Kermadec records, the distribution
is essentially Gondwanian.
of the genus in the combination of 4-segmented
ovigers and lacking auxiliary claws.
Etymology
This species is dedicated to Mr. David Staples, of
Melbourne, in recognition of his works on Austral-
ian Pycnogonida.
Biogeography
Child (1983: 699, 1988: 55) has suggested a ‘west-
ern Indian Ocean corridor’ and a ‘western Pacific
corridor’ to explain the presence of Austrodecus
species on Aldabra Atoll (Indian Ocean), and in
warm or warm-temperate waters in the western
Pacific (Kermadec Islands, Palau, Japan). Stock
(1957: 24, fig. 10) presumed a relationship to the
plate tectonics of the southern hemisphere land
masses, long before plate drift was a fashionable
subject. Fig. 9 shows that (when the distribution of
Austrodecus is plotted on a late Mesozoic map of
the continents), the genus is essentially Gondwan-
lan, with extensions along the western margin of
the Pacific Ocean.
REFERENCES
Child, C. A., 1983. Pycnogonida of the western Pacific
islands, II Guam and the Palau islands. - Proceedings
of the biological Society of Washington 96: 698-714.
Child, C. A., 1988. Pycnogonida from Aldabra atoll. — Bul-
letin of the biological Society of Washington 8: 45-78.
Clark, W. C., 1972a. Pycnogonida of the Snares Islands. —
New Zealand Journal of marine and freshwater Re-
search 5: 329-341.
Clark, W. C., 1972b. Pycnogonida of the Antipodes Islands.
— New Zealand Journal of marine and freshwater Re-
search 5: 427-452.
Gordon, I., 1944. Pycnogonida. - Report of the B.A.N.Z.
Antarctic Research Expedition (B) 5 (1): 1-72.
Hedgpeth, J. W. & J. C. McCain, 1971. A review of the
pycnogonid genus Pantopipetta (family Austrodeci-
dae, emended) with the description of a new species.
— Antarctic Research Series 17: 217-229.
Pushkin, A. F., 1977. New species of Austrodecidae from
the Antarctic. - Exploration of the Fauna of the Seas,
Academy of Sciences of the U.S.S.R. 21 (29): 54-59.
Smith, A. G., & J. C. Briden, 1977. Mesozoic and Cenozoic
paleocontinental maps: 1-63 (Cambridge University
Press).
Staples, D., 1977. Pycnogonida, the "nobodies”. - Austra-
lia’s Oceans (Journal of the Oceans Society of Austra-
lia) 1: 122-125.
Dj
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Stock, J. H., 1954. Pycnogonida from Indo-West Pacific,
Australian and New Zealand waters. - Videnskabelige
Meddelelser fra Dansk naturhistorisk Forening i
Kjabenhavn 116: 1-168.
Stock, J. H., 1957. The pycnogonid family Austrodecidae.
— Beaufortia 6: 1-81.
272
Stock, J. H., 1968. Pycnogonida collected by the Galathea
and Anton Bruun in the Indian and Pacific Oceans. -
Videnskabelige Meddelelser fra Dansk naturhistorisk
Forening i Kjobenhavn 131: 7-65.
J. van TOL
National Museum of Natural History, Leiden
ZOOLOGICAL EXPEDITIONS TO THE KRAKATAU
ISLANDS, 1984 AND 1985: ODONATA
Descriptions and records of Malesian Odonata, 2
Tol, J. van, 1990. Zoological expeditions to the Krakatau Islands, 1984 and 1985: Odonara.
Descriptions and records of Malesian Odonata, 2. - Tijdschrift voor Entomologie 133: 273-
279, figs. 1-11, tab. 1 [ISSN 0040-7496]. Published 14 December 1990.
A list of the Odonata collected by the members of the Krakatau Expeditions (1984 and 1985),
organized by the La Trobe University (Melbourne) and LIPI (Jakarta). Most of the species
mentioned are not from the Krakatau Islands, but from westernmost Java (Jawa) or
southernmost Sumatra (Sumatera). Agriocnemis pygmaea (Rambur) and Orthetrum t.
testaceum (Burmeister) are recorded from the Krakatau Islands for the first time.
J. van Tol, National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The
Netherlands.
Key words. - Odonata; Malesia; Krakatau; Java; Sumatra.
Members of most entomological expeditions to
the Krakataus have made collections of dragonflies
(Odonata). Yukawa & Yamane (1985) recently
summarized the present knowledge. Their list of
references should be completed with the work of
Ris (1912), since that publication provides the iden-
tifications of the specimens collected by Jacobson in
May 1908, and only referred to as ‘spec. 1° and ‘spec.
2’ by Jacobson (1909). Thornton & New (1988b)
gave some preliminary results of the present mate-
rial and some sight observations.
The present collection of Odonata includes con-
siderably more species, mainly due to sampling ac-
tivities in the southernmost part of Sumatra and the
westernmost part of Java. A detailed account of the
general results of these expeditions were given by
Thornton and Rosengren (1988), while Thornton
& New (1988a) deal with the freshwater commun-
ities. Presently, the only known relatively perman-
ent water bodies on the Krakatau Islands are con-
crete-lined ponds, one on Sertung and two on
northern Panjang. Before 1940 there were several
pools and a brackish-water lake with breeding pop-
ulations of eurytopic Odonata, e.g. Pseudagrion mi-
crocephalum (Rambur), Ischnura senegalensis
(Rambur), Diplacodes trivialis (Rambur) and Pan-
tala flavescens (Fabricius) (Dammerman 1948).
The following systematic account lists all species
collected during the expeditions of 1984 and 1985.
Species collected on the Krakatau islands Rakata,
Sertung, Anak Krakatau or Panjang are marked
with an asterisk. All records are given in full, but
data on longitude-latitude are summarized in the
next paragraph.
LOCALITIES
Sumatra. - Liwa (1), sweeping, 5°04’S 104° 03’E;
Liwa (2), secondary forest, sweeping, 700 m, 5° 04'S
104° 03’E; Liwa (4), 5°04’S 104° 03’E.
Krakatau Islands. — Rakata, Owl Bay, 6°09'S
105°28'E; Panjang, 6°05’S 105°28'E; Anak Kraka-
tau, sweeping, 6°06'S 105°26'E; Sertung, forest,
6°05’S 105°23’E; Sertung, Spit, 6°04’S 105°24’-
DONE:
Java. - Ujung Kulon, Pulau Peucang, 6°45’S
105° 15’E; Ujung Kulon, Cideon, 6°46'S 105° 15’E;
Ujung Kulon, Cibunar, 6° 48'S 105° 17’E; Hills be-
hind Carita.
The present and former names of the islands are:
Rakata, also known as Rakata Besar and Krakatau;
Panjang, also known as Rakata Kecil and Lang
eiland; Sertung, also known as Verlaten eiland.
Compilation of information on topography and the
history of events are to be found in Dammerman
(1948), and the more recently published works of
Simkin & Fiske (1983) and Thornton & Rosengren
(1988).
SYSTEMATIC LIST
The systematic list follows Davies & Tobin
(1984, 1985). Records for each species are arranged
in chronological order. Short comments on distri-
bution and abundance of the species are given, us-
275
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
ually based on Lieftinck's publications (Lieftinck
1934, 1935, 1953 and 1954) and the RMNH collec-
tion, Leiden. Special attention was paid to notes
giving details about occurrence in southern Sumatra
and western Java. Especially Lieftinck’s records of
‘Mt Tanggamoes’ [= Gunung Tanggamus] are of
great interest for the interpretation of the observa-
tions from Liwa. The status of Odonata before 1940
is well established through many collecting trips of
entomologists working for the Bogor museum (for-
merly Buitenzorg).
Abbreviations used: RMNH = National Mu-
seum of Natural History, Leiden.
Family Amphipterygidae
Devadatta argyoides (Selys, 1859)
Sumatra, Liwa (2), 191 BB, 1 Sep 1984, 1 female.
Although generally uncommon, it was very
abundant in the Gn Tanggamus area in December
1934 (Lieftinck 1935).
Family Calopterygidae
Neurobasis chinensis chinensis (Linnaeus, 1758)
(fig. 1)
Sumatra, Liwa (2), 191 BD, 1 Sep 1984, 1 male;
Idem, 191-BA, 1 female.
The nominal subspecies is common on Sumatra,
including the southernmost part. Javanese popula-
tions are ranked as a separate subspecies, N. chinen-
sis florida Hagen.
Vestalis luctuosa (Burmeister, 1839)
Sumatra, Liwa (2), 191 BC, 1 male.
Java, Ujung Kulon, Cidaon, 190 GA and 190 HC,
15 Sep 1984, 1 male, 1 female; Idem, 190 ND, 16
Sep 1984, 1 male.
Vestalis luctuosa is a common species on Java, but
on Sumatra it is confined to the southernmost part.
V. amoena Selys, a common species of this genus on
Sumatra, was not represented.
Females of Newrobasis chinensis and the re-
gional Vestalis species are superficially very similar,
and are not easy to separate in the field, especially
so, since in many specimens of N. chinensis the
pseudopterostigma is absent. Both genera are most
easily distinguishable based on their wing venation.
The Arculus is oblique, but not angulated in
Vestalis, while it is conspicuously angulated in New-
robasis (figs. 1 and 2) (Fraser 1933). Besides, Neu-
robasis has cross-veins in the median (basal) space.
Both Vestalis species can be identified on the col-
oration of the labium (cf. Schmidt 1934: figs. 19a-
b).
Vestalis lugens Selys
Sumatra, Liwa (2), 190 BE, 1 Sep 1984, 2 females;
Idem, Liwa (1), 191 AA, 6 Sep 1984, 1 female; Idem,
274
Figs. 1-2. Wing base in Calopterygidae. - 1 (above), Ves-
talis amoena; 2, (below) Neurobasis chinensis.
Liwa (2), 191 AB, 7 Sep 1984, 1 female.
See remarks under V. /uctuosa.
Family Chlorocyphidae
Rhinocypha angusta (Selys, 1853)
Sumatra, Liwa (2), 191 BD, 1 Sep 1984, 1 male.
Common on Sumatra, but absent from Java.
Rhinocypha selysi Kriiger, 1898
Sumatra, Liwa (2), 191 BC, 1 Sep 1984, 1 male.
Species confined to Sumatra (contra Lieftinck
1934, cf. Lieftinck 1954). Generally a rare species,
except in southern Sumatra. In the RMNH collec-
tion 18 males and 13 females are represented. Most
specimens are from the Gn Tanggamus area and
Benkulen; the other sites are (from north to south):
Deli, Bolimbingan, 600 m (leg. Straatman) [locality
not found]; Emmahaven, Padang [1°00'S
100° 22’E] (leg. Van der Starre); Sandaran Agung
(Kurintji) [= probably Sanggaran agung, Kerintji;
2°08'S 101°31’E] (leg. Jacobson).
Family Euphaeidae
Euphaea variegata (Rambur, 1842)
Sumatra, Liwa (2), 191 BA, 1 Sep 1984, 1 male.
Java, Ujung Kulon, Cidaon, 190 HC, 15 Sep 1984,
1 female.
Widespread and common on Sumatra and Java;
recorded as ‘very common’ from the S. Lampoeng
Residency by Lieftinck (1935). It was also found on
Panaitan Island (Lieftinck 1953).
Family Coenagrionidae
Agriocnemis femina (Brauer, 1868) (figs. 9-11)
Sumatra, Liwa (1), 191 AA, 6 Sep 1984, 1 female.
Eurytopic species, very common and widespread
from Bengal, Assam (Fraser 1933) eastwards to
northern Australia. See further notes under A. pyg-
maed.
* Agriocnemis pygmaea (Rambur, 1842)
(figs. 5-8)
Krakatau Is. - Anak Krakatau, 215.DE, 21 Aug
1985, 1 female [short-lobed form, headless |.
New to the Krakataus. Habitat: all kinds of stag-
nant waters. Presumably a wind-borne species
(Lieftinck 1954). Very widespread , and usually very
common in the afrotropical, oriental and australian
region.
VAN Tor: Odonata Krakataus 1984-1985
The females of Agriocnemis are not easy to dis-
tinguish, since both A. femina and A. pygmaea are
polytypic. In this area also another species, A. min-
ima Selys (figs. 3-4), has to be taken into account.
I have now provisionally examined material avail-
able in the RMNH collection (figs. 3-11). It seems
that at least two forms of both species based on the
shape of the prothorax, can be recognized. Females
of A. pygmaea may have a strongly developed hind-
lobe with a median depression, or the hindlobe is
nearly completely lacking (figs. 5-8). Apart from
these structural forms, there also exist two colour
forms, an orange-red and a greenish-brown form.
Therefore, as already stressed by Ris (1916), four
forms can be recognized, since differently shaped
hindlobes of the pronotum occur in both colour
forms.
According to Ris (1916), Agriocnemis femina
10 11
Figs. 3-11. Prothorax in females of Agriocnemis, left lateral and dorsal view. — 3-4, A. minima, Java; 5-6, A. pygmaea
long-lobed form from Sulawesi; 7-8, À. pygmaea, short-lobed form from Ceylon; 9, A. femina, form with medially
depressed hind lobe, northern Borneo; 10-11, A. femina, form with upright hind lobe, Sumatra.
275
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
falls apart in at least four structurally different
forms. Also the males of this species show geogra-
phical variation. Females of A. femina always have
a well-developed hindlobe, but it is strongly upright
in specimens from Ceylon, or it is conspicuously
depressed in the middle in more eastern popula-
tions (figs. 9-11). According to Lieftinck (1954: 72
footnote) two well definable subspecies occur on
Sumatra.
Family Platycnemididae
Coeliccia membranipes membranipes (Rambur,
1842)
Sumatra, Liwa (1), 191 AA, 6 Sep 1984, 1 female.
Common species on Java and Sumatra in slowly
flowing waters.
Family Aeshnidae
Gynacantha basiguttata Selys, 1882
Java, Ujung Kulon, 190 AG, 17 Sep 1984, 1 male.
Considered as ‘rare’ on Java by Lieftinck (1934);
more common on Sumatra, and also known from
Billiton, Simalur, Enggano and western Borneo
(collection RMNH). Apparently not rare on Panai-
tan Ísland (Lieftinck 1953). Breeds in forest pools.
The adults fly only at dusk, as all Gynacantha
species.
Family Libellulidae
Agrionoptera insignis insignis (Rambur, 1842)
Java, Ujung Kulon, Pulau Peucang, 15 July 1984,
190 AD, 1 male; Idem, 190 AF, AK, AP, AR, Sep
1984, 1 male 3 females; Idem, 190 AC, 20 Sep 1984,
1 female; Java, Ujung Kulon, Cibunar, 190 GB, Sep
1984, 1 male; Java, Ujung Kulon, P. Peucang, 24.1,
22 Sep 1984, 1 female.
Confined to the coastal area of a significant part
of the oriental region. Locally abundant in e.g. man-
grove forests. It breeds in shady pools in forests
(Lieftinck 1954). Although it was once ‘one of the
commonest, and certainly the most striking’ (Lief-
tinck 1934) dragonfly of the Krakataus, it was not
collected there during this expedition, nor by Yu-
kawa & Yamane (1985). The collection made in
1908 by Jacobson (Ris 1912) consisted of this spe-
cies and Diplacodes trivialis (Rambur, 1842) only.
Cratilla lineata assidua (Lieftinck, 1953)
Java, Ujung Kulon, Pulau Peucang, 11 Sep 1984,
1 male.
A woodland species, obviously uncommon on
Java. The nominotypical subspecies occurs widely
on Sumatra and was once collected on the Krakatau
islands, while subspecies assidua Lieftinck was re-
corded from Panaitan Island (Lieftinck 1953).
276
Lathrecista asiatica (Fabricius, 1798)
Java, Ujung Kulon, Pulau Peucang, 190 AJ, 13
Sep 1984, 1 male.
Although widespread in the oriental region, Lief-
tinck (1954) considered it uncommon on Java.
Lyriothemis magnificata (Selys, 1878)
Java, Ujung Kulon, Pulau Peucang, 190 AQ, 3 Sep
1984, 1 female.
A rare species from Sumatra, Billiton and Java.
Unknown from western Java according to Lieftinck
(1934), but in the RMNH collection several spec-
imens from this area are available, e.g. Udjung
Kulon, Tjibunar [river], 28 Nov 1958, 1 female.
Orthetrum sabina (Drury, 1770)
Sumatra, Liwa (1), white light, 190 AC, 31 Aug
1984, 1 female; Idem, Liwa (2), 191 BE, 1 Sep 1984,
1 male.
Java, Ujung Kulon, Pulau Peucang, 190 AN, 17
Sep 1984, 1 male; Idem, Carita, 310 A, 11 Aug 1985,
1 female. Widespread and common species; abund-
ant in cultivated areas. Collected on the Krakataus
by Yukawa & Yamane (1985).
* Orthetrum testaceum testaceum (Burmeister,
1839)
Sumatra, Liwa (2), 191 BA and 191 BC, 1 Sep
1984, 2 males; Sumatra, Liwa (4), 191 DA, 7 Sep
1984, 1 male.
Krakatau Is., Penjang, 3 Sep 1984, 192 BA, 1
male.
New to the Krakatau Islands. Common in the
lowlands of Java and Sumatra, and also known from
Panaitan Island (Lieftinck 1953). Breeding in all
kind of waters (Lieftinck 1934).
* Diplacodes trivialis (Rambur, 1842)
Sumatra, Liwa (1), 191 AA, 6 Sep 1984, 1 female.
Krakatau Is, Anak Krakatau, 192 AA, 13 Sep
1984, 1 female; Krakatau Is., Anak Krakatau, 310B;
15 Aug 1985, 1 female; Idem, 310 C, 17 Aug 1985,
1 male; Krakatau Is., Rakata, Owl Bay, 310 F, G and
H, 26 Aug 1985, 1 male 2 females.
Very common and widespread species. Recorded
from the Krakataus since 1908.
Neurothemis fluctuans (Fabricius, 1793)
Sumatra, Liwa (4), 191 DB, 7 Sep 1984, 3 males
1 female; Idem, 191 DC, same date, 2 males.
See remarks under the following species.
Neurothemis fluctuans (Fabricius, 1793) / N.
ramburii (Brauer, 1866)
Java, Ujung Kulon, Pulau Peucang, 190 AQ, Sep
1984, 1 male; Idem, 190 AH, Sep 1984, 1 female
(homeochrome); Idem, (no number), 11 Sep 1984,
1 female (homeochrome); Idem, 190 AI, 11 Sep
1984, 1 female (homeochrome); Idem, 190 AL, 13
Sep 1984, 1 male; Idem, 190 AE, 15 Sep 1984, 1
female (heterochrome); Idem, 24.1, 22 Sep 1984, 1
female (heterochrome); Ujung Kulon, 190 GA, 20
Sep 1984, 1 female (heterochrome).
According to Lieftinck (1954) Newrothemis fluc-
tuans is rare and extremely local on Java, while
Neurothemis ramburt is widespread but very local
there.
I am unable to identify the specimens from
Ujung Kulon; the specimens from Sumatra seem to
belong to N. fluctuans. It is well known that iden-
tification of the genus Newrothemis meets many
problems in this part of their range. Lieftinck
(1954: 151 footnote) mentions hybridization be-
tween N. fluctuans, N. ramburi and N. terminata,
three species frequently flying on the same site.
Lieftinck (1953) records all three species from Pa-
naitan Island, and according to Lieftinck (1954)
they can usually be kept apart fairly easily. I have
examined large series in the RMNH collection, all
identified by Lieftinck, but relevant characters seem
to show significant variation between populations.
A further study of this complex is needed to judge
on the status of these taxa. Also Ris (1911: 567) has
expressed his doubts about the specific status of
these taxa. An extensive study of the variation of
the characters involved (e. g. size of wing marking,
number of cubito-anal cross-veins) could contribute
Table 1. Odonata on the Krakatau islands.
Rakata
Besar
Species Before Now
Coenagrionidae
Pseudagrion microcephalum
Ischnura senegalensis e
Agriocnemis femina
Agriocnemis pymaea
Libellulidae
Agrionoptera insignis e
Cratilla lineata e
Raphismia bispina
Acisoma panorpoides La
Orthetrum sabina
Orthetrum glaucum
Orthetrum testaceum
Diplacodes trivialis
Crocothemis servilia
Neurothemis terminata
Neurothemis tullia
Pantala flavescens
Rhyothemis phyllis
Tramea limbata euryale
Macrodiplax cora
Aeshnidae
Anax guttatus
Total number of species
Rakata
Kecil
Before
Fi
VAN Tor: Odonata Krakataus 1984-1985
to our understanding of the historical biogeography
of this species complex.
Neurothemis terminata Ris, 1911
Java, Ujung Kulon, hills behind Carita, 11 Aug
1985, 1 male; Idem, Cidaon, 190 HB, 15 Sep 1984,
1 male.
A common species on Java. See also remarks
under N. fluctuans / N. ramburii.
* Neurothemis tullia feralis (Burmeister, 1839)
Krakatau Is., Anak Krakatau, outer rim, Malaise
trap 21-24 Aug 1985, 213 EA, 1 male.
Second species known from Anak Krakatau, and
only the second record of this species from the
Krakataus (first record: Rakata, 23 Oct 1923). It is
rather rare and local on Java (Lieftinck 1934); there
are no reliable records from Sumatra. It inhabits
marshlands and weedy ponds.
* Rhyothemis phyllis phyllis (Sulzer, 1776)
Krakatau Is., Panjang, 192 BB, 3 Sep 1984, 1
male.
Rather common species on Java, probably also
breeding in oligohaline waters. The present record
is the first of this species from Panjang, and only
the second record for the Krakatau islands (first
record: Rakata, 1921).
Anak
Krakatau
Sertung
Now Before Now Before Now
No
—
QW
—
—
Ww
ZT
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
Camacinia gigantea (Brauer, 1867)
Java, Ujung Kulon, Pulau Peucang, 190 AM, 18
Sep 1984, 1 female.
Widespread from Burma and Assam eastwards
to Eastern New Guinea. Usually rather uncommon,
restricted to low altitudes in non-cultivated areas.
* Pantala flavescens (Fabricius, 1798)
Krakatau Is., Sertung Spit, at camp, 310 D, 18
Aug 1985, 1 male; Idem, Sertung forest, 310 E, 19
Aug 1985, 1 male.
A very common circumtropical species and
strong flier.
DISCUSSION
The dragonfly fauna of the Krakataus is only a
poor representation of the fauna of the ‘mainlands’
of Java and Sumatra. This can even be demonstrated
with the present data, but especially when all data
available about the fauna of Java and Sumatra, or
even those about that of Panaitan Island, are com-
pared with those from the Krakataus.
The reason for the high turnover rate of the
species involved must be the fact that there is hardly
any water available as breeding site for freshwater
invertebrates. Most specimens collected should be
considered as stragglers from Java or Sumatra, and
must have reached the Krakataus by active flight or
transported by the wind. The fauna of Panaitan
Island (c. 150 km?, c. 10 km from the mainland of
Java) was found to be far more mature in 1951 and
also included several stream-dwelling species. Lief-
tinck (1953) concluded that the dragonfly fauna of
that island was not entirely composed of species
that had reached the island accidentally, but that
certain species must have inhabited it since the time
it was a part of Java. Not all species found on the
Krakatau islands (table 1) were also collected on
Sumatra and Java by the present expeditions. How-
ever, they are all are eurytopic or inhabiting coastal
biotopes, where they are common, and usually also
abundant, in southern Sumatra and/or western Java
(based on Lieftinck 1954).
Nevertheless, in future one may expect an indi-
genous dragonfly fauna on the Krakataus when the
forests have become more mature. Several highly
specialized species of Zygoptera as well as Anisop-
tera, are known as breeders in phytotelmata, e.g.
treeholes of decaying logs, bamboo stumps, or wa-
terholes at the leaf-base of Pandanus (cf. Corbet
1983). Examples of this fauna are the members of
the libelluline genus Lyriothemis Brauer (cf. Lief-
tinck 1954, Kitching 1986), which breed in water-
filled treeholes, and the pseudagrionine genus Am-
phicnemis Selys, supposed to breed in water at the
leaf-bases of Pandanus trees. This kind of Odonata
may find a breeding site on the Krakataus in due
time. It may even be true that representatives of
278
this kind of dragonflies already inhabit the Kraka-
taus, since most of them are very hard to collect by
their inconspicuous behaviour.
From a zoologeographical point of view it is of
interest to mention that the fauna of the Krakataus
seems to be Sumatran rather than Javan, contrary
to the Panaitan Island.
ACKNOWLEDGEMENT
I am indebted to Professor I. W. B. Thornton
(Melbourne, Australia) for entrusting this collec-
tion to me.
REFERENCES
Corbet, P. S., 1983. Odonata in Phytotelmata. In
Phytotelmata: Terrestrial plants as hosts for aquatic
insect communities (eds. J.H. Frank & L.P. Lounibos).
pp. 29-54. Marlton, New Jersey: Plexus.
Dammerman, K. W., 1948. The fauna of Krakatau 1883-
1933. Verhandelingen der Koninklijke Nederlandsche
Akademie van Wetenschappen, afd. Natuurkunde
(Section 2) 44: i-xii + 1-594 + plate 1-11.
Davies, D. A. L. & P. Tobin, 1984. The dragonflies of the
world. A systematic list of the extant species of Odo-
nata. Volume 1. Zygoptera, Anisozygoptera. - Rapid
Communications Societas Internationalis Odonatolog-
ica, Supplement 3: 1-127.
Davies, D. A. L. & P. Tobin, 1985. The dragonflies of the
world. A systematic list of the extant species of Odo-
nata. Volume 2. Anisoptera. - Rapid Communications
Societas Internationalis Odonatologica, Supplement 5:
1-151.
Fraser, F.C. 1933 Fauna of British India including Ceylon
and Burma. Odonata. Volume 1. Taylor & Francis,
London.
Jacobson, E. R., 1909. De opneming van de Krakatau-
groep in Mei 1908. 5. De nieuwe fauna van Krakatau.
— Jaarverslag van den Topographischen dienst in Ne-
derlandsch-Indié 1908: 192-206 (sep. 43-57), 1 table
excl.
Kitching, R. L., 1986. A dendrolimnetic dragonfly from
Sulawesi (Anisoptera: Libellulidae). - Odonatologica
15: 203-209.
Lieftinck, M. A., 1934. An annotated list of the Odonata
of Java, with notes on their distribution, habits and life-
history. - Treubia 14: 377-462 + 1 plate.
Lieftinck, M. A., 1935. A synopsis of the Odonata (Dra-
gonflies) of Sumatra. - Miscellanea Zoologica Suma-
trana 92-93: 1-23.
Lieftinck, M. A., 1953. Notes on some Odonata from
Panaitan Island, Sunda Strait (West Java). - Idea 9: 51-
56, 2 maps.
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.
Ris, F, 1911. Libellulinen 5. - Collections Zoologiques du
Baron Edm. de Selys Longchamps 13: 529-700 + plate
V.
Ris, FE, 1912. Uber Odonaten von Java und Krakatau
gesammelt von Edward Jacobson. - Tijdschrift voor
Entomologie 55: 157-183 + plate 6-8.
Ris, F, 1916. H. Sauter’s Formosa-Ausbeute. Odonata.
(Mit Notizen über andere ostasiatische Odonaten). —
Supplementa Entomologica 5: 1-81, figs. 1-47, plates
1-3 excl.
Schmidt, E., 1934. Odonata der Deutschen Limnologi-
schen Sunda-Expedition. I. Imagines. Mit Beschrei-
bungen zweier neuer Ictinus aus Celebes und Neu-
Guinea. — Archiv für Hydrobiologie, Suppl. 13: 316-
397 figs. 1-93, tabs 1-3, tabs 1 + Plate 14-17 excl.
Simkin, T. & R. S. Fiske, 1983. Krakatau 1883. The vol-
canic eruption and its effects. - Smithsonian Institu-
tion Press, Washington DC: 1-464, figs. 1-139, plate 1-
16 excl.
Thornton, I. W. B. & T. R. New, 1988a. Freshwater com-
munities on the Krakatau Islands. — Philosophical
Transactions of the Royal Society of London (Series B)
322: 487-492, fig. 1, tab. 1.
VAN ToL: Odonata Krakataus 1984-1985
Thornton, I. W. B. & T. R. New, 1988b. Krakatau inver-
tebrates: the 1980s fauna in the context of a century of
recolonization. — Philosophical Transactions of the
Royal Society of London (Series B) 322: 493-522, figs.
1-2, tabs. 1-10.
Thornton, I. W. B. & N. J. Rosengren, 1988. Zoological
expeditions to the Krakatau Islands, 1984 and 1985:
General introduction. — Philosophical Transactions of
the Royal Society of London (Series B) 322: 273-316,
figs. 1-24, tabs. 1-10, plates 1-14 excl.
Yukawa, J. & Yamane, S. 1985. Odonata and Hemiptera
collected from the Krakataus and surrounding islands,
Indonesia. - Konty 53: 690-698.
Received: 10 September 1990
Accepted: 27 September 1990
279
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 133, 1990
280
NEW TAXA DESCRIBED IN VOLUME 133
ACARI
ASGCELOUChAMPÈLE A SANYEO EEE IT 247
Ascetolichus microthrix Pérez & Atyeo ..... 249
Ascetolichus ruidus Pérez & Atyeo ......... 249
COLEOPTERA
Carphurus albipennis Wittmer ............ 109
Carphurus basilimbatus Wittmer .......... 112
Carphurus brendelli Wittmer ............. 113
Carphurus dumogaensis Wittmer .......... 110
Carphurusilyale Watters Seal: 1415
Carphurus pallidicornis Wittmer .......... 109
Carphurus partepallidus Wittmer ......... 112
Garphurus storks Wittmenes u. aan 114
Carphurus strigilatus Wittmer ............ 111
Carphurus testaceolimbatus Wittmer ....... 113
Carphurus totopallidus Wittmer .......... 111
Dromanthomorphus ranuensis Wittmer .... 118
Falsolaius semicostatus Wittmer .......... 177
Falsolaius sulawesiensis Wittmer .......... 116
Bamıssdeniicollsı Wattmer reren ee 119
EON IEN SMN IEEE GRAN AR 118
AUST ANUER ISA IEEE PETE EE: 119
Paracarphurus sulawesiensis Wittmer ...... 115
Telocarphurus nebulosus Wittmer ......... 115
COLLEMBOLA
Superodontella ciconia Bedos &
DETAEVEN caste. un Myc tit. rere erry tt EI 1)
Superodontella flammata Bedos &
lWeharyeng rn Re EER selon eee 25
Superodontella gouzei Bedos & Deharveng 21
Superodontella longispina Bedos & Deharveng
Bean. dan aduk aye te 21
DIPTERA
Cryptochironomus judicius Chaudhuri & Chatto-
padhyayse AE ME AE AVE RO RAD De DI 154
Fletcherimyia abdita Pape ............... 48
Harnischia tenuitubercula Chaudhuri & Chattopad-
AMENER 20 Manag LOA AEN Ien o 158
Microcerella adelphe Pape ............... 52
Microcerella aulacophyto Pape ............ 49
(nom. nov. for Aulacophyto auromaculata
Townsend)
Microcerella austrohartigia Pape .......... 51
(nom. nov. for Austrohartigia bicoloricauda
Lopes)
Microcerella bermuda Pape .............. DIL
WMicnocenellalboeta Papel. 22: 222.2... 50
(nom. nov. for Boettia ecwatoriana Lopes)
Microcerella boettcherimima Pape ......... 49
(nom. nov. for Boettcherimima hypopygialis
Lopes)
Microcerellaicanchiabape eae ee. 49
(nom. nov. for Carchia andina Lopes)
Macnocercllagujuytapapem ren. EEE 49
(nom. nov. for Jujuyia alvarengai Lopes)
Paracladopelma aratra Chaudhuri &
Chattopadhyay 4/404 ees ore 160
Paracladopelma sacculifera Chaudhuri &
Chattopadhyay sunt ne Cee ee eee 161
Polypedilum circulum Chaudhuri &
Chattopadhyay ines er PR eee ee 162
Sarcophaga (Discachaeta) amputata Pape .... 39
Sarcophaga (Pierretia) inlicida pape ........ 41
Spirobolomyia latissima Pape ............. 54
Tripanurga guatemala Pape .............. 57
LEPIDOPTERA
Biselachista spinigera Sruoga ............. 79
Cosmiotes kopetdagica Sruoga ............ 83
Cosmsotenpallenworuoga Garena TIE 83
Deltas bobaga van Mastrigt .............. 200
Deltas catocausta eeft van Mastrigt ........ 201
Delias luctuosa kuning van Mastrigt ....... 202
Deltas nieuwenhuisi van Mastrigt ......... 202
(nom. nov. for Deltas leucias roepkei Nieuwen-
huis & Howart)
Deltas nieuwenhuisi poponga van Mastrigt 204
Deltas oktanglap van Mastrigt ............ 201
Delrastsiert var Mas trict =. Sun u GSC 200
DelasisinakvantMastrigth seater e 200
Delias wollastoni abmisibilensis van Mastrigt 202
Elachista fuscofrontella Sruoga ............ 76
Elachista megagnathos Sruoga ............ 79
Elachista multipunctata Sruoga ........... 75)
Euchromius cornus Schouten ............. 265
RenttiabilobaSruogan OICR TI ANO 76
Stigmella rolandi van Nieukerken ......... 239
Trifurcula austriaca van Nieukerken ....... 245
Trifurcula coronillae van Nieukerken ....... 21197,
Trifurcula iberica van Nieukerken ......... 228
Trifurcula josefklimeschi van Nieukerken 225
Trifurcula luteola van Nieukerken ......... 215
Trifurcula puplesisi van Nieukerken ....... 215
Trifurcula silviae van Nieukerken ......... 229
Trifurcula victoris van Nieukerken ......... 220
ODONATA
Leptogomphus pasta van Tol ............. 97
BhylioeyelalhamaraaBelle Era 29
Progomphus gel NBE 27
PYCNOGONIDA
Austmnodecus staples Stock 2... Hann ee 269
Vv
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CORRIGENDA
The figures 72-73 of the paper by J. Belle, A revision of the New World genus Newraeschna Hagen, 1867
(Odonata: Aeshnidae) published in volume 132 (2), page 283, unfortunately are depicted too large. The
reproductions below show these figures in the correct size.
Figs. 72-73. — Neuraeschna titania spec. nov., & holotype; 73, Neuraeschna titania spec. nov., 9 paratype from same
locality as holotype.
vili
Tijdschrift voor Entomologie
Volume 133, no. 2
Articles
121 B. Aukema
Taxonomy, life history and distribution of three closely related species of the
genus Calathus (Coleoptera: Carabidae).
143 J. Belle & J. van Tol
Anomalagrion hastatum (Say), an American damselfly indigenous to the Azores
(Odonata, Coenagrionidae).
149 P. K. Chaudhuri & S. Chattopadhyay
Chironomids of the rice paddy areas of West Bengal, India (Diptera:
Chironomidae).
197 H. J. G. van Mastrigt
New (sub)species of Delias from the central mountain range of Irian Jaya
(Lepidoptera: Pieridae).
205 E. J. van Nieukerken
The Trifurcula subnitidella group (Lepidoptera: Nepticulidae): taxonomy,
distribution and biology.
239 E. J. van Nieukerken
Stigmella rolandi sp. n.: a widespread southern European species on Rosa
(Lepidoptera: Nepticulidae)
245 T.M. Pérez & W. T. Atyeo
New taxa of feather mites (Acarina, Pterolichidae) from Megapodes (Aves,
Megapodiidae).
251 D.L. J. Quicke & C. van Achterberg
The type specimens of Enderlein's Braconinae (Hymenoptera: Braconidae)
housed in Warsaw.
265 R. T. A. Schouten
A new species of Euchromius Guenée, 1845 (Lepidoptera: Pyralidae:
Crambinae) from Australia.
269 J. H. Stock
A new species of Austrodecus (Pycnogonida) from New South Wales, Australia.
273 J. van Tol
Zoological expeditions to the Krakatau Islands, 1984 and 1985: Odonata.
Descriptions and records of Malesian Odonata, 2.
Book announcements and reviews
268 T.B. Larsen, The butterflies of Egypt; O. |. Merzheevskaya, Larvae of owlet
moths (Noctuidae); P. W. Oman, W. J. Knight & M. W. Nielson, Leafhoppers
(Cicadellidae): a bibliography, generic check-list and index to the world
literature 1956-1985; D. |. Williams & G. W. Watson, The scale insects of the
Tropical South Pacific region, 3.
© Nederlandse Entomologische Vereniging, Amsterdam
Published 14 December 1990 ISSN 0040-7496
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