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Volume 137, no. I, 1994 ISSN 0040-7496
Tijdschrift
\Svoor
— —Entomologie
A journal of systematic and evolutionary |
entomology since 1858
?
d: 2)
Tg N EIS
Published by the Nederlandse Entomologische Vereniging
Tijdschrift voor Entomologie
A journal of systematic and evolutionary entomology since 1858
Scope
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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
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Usually, such papers will only be published when space allows.
Editors
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Co-editors
A. W. M. Mol (1990) and R. T. A. Schouten (1990)
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The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the
‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society),
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Instructions to authors
Published with index of volume 136 (1993).
Graphic design
Ontwerpers B.V., Aad Derwort, ’s-Gravenhage
A. J. DE BOER
Institute for Systematics and Population Biology (Zoölogisch Museum), University of Amsterdam
THE TAXONOMY AND BIOGEOGRAPHY OF THE
LORIAE GROUP OF THE GENUS BAETURIA STAL,
1866 (HOMOPTERA, TIBICINIDAE)
Boer, A. J. de, 1993. The taxonomy and biogeography of the loriae group of the genus Baeturia
Stal, 1866 (Homoptera, Tibicinidae). — Tijdschrift voor Entomologie 137: 1-26, figs. 1-102,
table 1. [ISSN 0040-7496]. Published 15 July 1994
The loriae group is proposed for a presumably monophyletic group of 9 species. Species of the
group combine characters seemingly synapomorphic for the species of Baeturia Stal, 1866, with
characters seemingly synapomorphic for the species of Gymnotympana Stal, 1861. Though the
generic relationships of the /oriae group are not clear, it is included here in the genus Baeturia,
on account of shared male genital characters. One species (B. loriae Distant, 1897) is rede-
scribed and eight species (B. bemmeleni, B. daviesi, B. fortuini, B. hamiltoni, B. hartonoi, B. pi-
grami, B. silveri, and B. wegeneri) are described as new. B. loriae is transferred back from
Gymnotympana to Baeturia. A key to the males is presented. A redescription of B. tenuispina
Blöte, 1960 is included in this publication, since a character of the clasper indicates a relation-
ship with some species of the loriae group. Nevertheless, the attribution of that species to the
loriae group is far from certain. The loriae group is endemic to New Guinea, most species are
restricted to the central and eastern mountain ranges of that island.
A. J. de Boer, Institute for Systematics and Population Biology (Zoölogisch Museum),
University of Amsterdam, PO Box 4766, 1009 AT Amsterdam, The Netherlands.
Key words. — Baeturia, loriae group, taxonomy, biogeography, New Guinea.
Current studies of the tibicinid cicadas of eastern
Malesia show, that a number of New Guinean and
Australian genera form a monophyletic group: ‘the
Baeturia and related genera complex’ (de Boer 1990,
1991). The largest genus of this group, Baeturia Stal,
1866, appears to be non-monophyletic in its present
concept, but, within that genus, several monophylet-
ic species groups have been recognized. Taxonomic
and biogeographic revisions have been published so
far for the B. nasuta group (de Boer 1982), the B. con-
viva group (de Boer 1986), the B. bloetei group (de
Boer 1989) and the B. viridis group (de Boer 1992).
These pages present the revision of a presumably
monophyletic group of nine species, for which the
name Baeturia loriae group is proposed. This group is
of particular interest from a taxonomic point of view,
since its species combine several characters regarded
apomorphic for Baeturia, with characters apomor-
phic for Gymnotympana Stal, 1861. The distribution
of these characters within the group is such, that any
reconstruction of the in-group relationships will un-
avoidably show several conflicting characters. The B.
loriae group is endemic to northern, eastern and cen-
tral New Guinea.
B. tenuispina Blôte has a character of the clasper in
common with some species of the B. loriae group, but
differs in most other respects. This species is not at-
tributed to the B. loriae group, but described here,
since it seems more related to this group than to any
of the other species groups of Baeturia.
MATERIAL AND METHODS
The material used for this study comes from the
following institutions:
AMS — Australian Museum, Sydney; BMNH
— Natural History Museum [formerly: British
Museum (Natural History)], London; BPBM —
Bernice P. Bishop Museum, Honolulu; CSIRO —
Commonwealth Scientific and Industrial Research
Organisation, Australian National Insects collection,
Canberra; KBIN — Koninklijk Belgisch Instituut voor
Natuurwetenschappen, Brussel; MSNG — Museo
Civico di Storia Naturale ‘G. Doria’, Genova; MVM —
Museum of Victoria, Melbourne; RMNH — Nationaal
Natuurhistorisch Museum (formerly: Rijksmuseum
van Natuurlijke Historie), Leiden; SMN — Staatliches
Museum fiir Naturkunde, Stuttgart; ZMA — Institute
for Systematics and Population Biology (Zoölogisch
Museum), Amsterdam.
The following sources have been used for tracing
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
the localities: “Atlas van tropisch Nederland’ (1938),
‘The Times Atlas of the World (Comprehensive
Edition)’ (1968), ‘Papua New Guinea a travel survi-
val kit by Tony Wheeler, 1988, Lonely Planet
Publications, Victoria, Australia and a ‘List of New
Guinea localities’ published by the Bernice P. Bishop
Museum (1966).
To examine the male genitalia, the pygofer was
pulled out, after overnight softening, with a sharp
needle inserted between the pygofer and the 8th ab-
dominal segment. The aedeagus was pulled out at the
same time, by inserting the needle between the clasp-
ers. Body and tegmen lengths of all specimens were
measured, other measurements are based on a maxi-
mum of ten specimens, when available.
PHYLOGENY
Classification of the B. loriae group
The generic classification of the loriae group pre-
sents a major problem. As stated above, the species
combine characters of Baeturia with characters of
Gymnotympana. Some of these characters appear to
be unique for either Baeturia or Gymnotympana (or
part of these genera). The distribution of these char-
acters within the loriae group is not homogeneous;
some species have more in common with Baeturia,
others more closely resemble Gymnotympana.
The following characters are found in the loriae
group and indicate a relationship with Baeturia, the
first four are presumed apomorphies for that genus.
(1) a strongly curved aedeagus, tapering to its apex,
with bluntly rounded lateral lobes and oval aede-
agus pore. This aedeagus shape is unique for
Baeturia.
(2) a narrow vertex, with large ocelli close together
(fig. 3). Two species of Gymnotympana have an
even narrower vertex. In other species of the
complex, the vertex is broader than in Baeturia.
(3) a broad and triangular middle spine on fore fe-
mur (fig. 19) is shared by all species of Baeturia.
(4) a fairly narrow and frontally bluntly rounded
postclypeus, distinctly protruding beyond vertex
lobes and, in most species, distinctly and angu-
larly swollen ventrally (figs. 3-4). A similar swel-
ling only occurs in some species of Baeturia, the
narrow, protruding, and rounded shape of the
postclypeus is possibly apomorphic for Baeturia.
(5) a rather stout caudodorsal beak, often angularly
bent and curved over anal valves. Other species
of the complex have a much shorter and erect
beak. The curved caudodorsal beak is probably
synapomorphic for several of the Baeturia spe-
cies groups together.
(6) almost straight parallel claspers, separated at the
base, and with a broad and rounded ventral hol-
low in the apical part. Such separated claspers are
unique for Baeturia and Scottotympana de Boer,
1991
(7) a distinct and angular clasper heel (in B. bemme-
leni, B. hamiltoni, and B. wegeneri). A clasper
heel is found in many species of Baeturia, but
not in other genera.
(8) rows of setae on veins of tegmina and wings (in
B. hamiltoni and B. wegeneri). Similar rows of
setae are found in many Baeturia species, but not
in any of the related genera. However, such setae
also occur, though more distinctly, in several ge-
nera of the Prasiini, a possible sister group of the
‘Baeturia and related genera complex’.
(9) dense brown speckling on body (in B. daviesi
and B. wegeneri). Such speckling, though strong-
ly variable in density, only occurs within
Baeturia.
(10) irregularly shaped brown spots on tegmina (in B.
pigrami and B. silveri, fig. 59). Such spots are on-
ly found in some species of Baeturia (e.g. B. gut-
tulinervis, B. guttulipennis and B. nasuta).
(11) a distinct crest between the aedeagal lobes. A si-
milar crest occurs in some species of the B. con-
viva group and some species related to that
group.
Other characters found in the B. loriae group do
not occur in other species groups of Baeturia, but
seem to indicate a relationship with Gymnotympana.
The first two must possibly be regarded as apomorp-
hies for Gymnotympana, others are either restricted to
part of that genus, or occur in several other genera as
well.
(1) sexual dimorphism in wing venation (males with
very slender anal fields, compare figs. 59 and 84)
is shared by all species of Gymnotympana, but
does not occur in other genera of the complex.
(Similarly slender anal fields are recorded for
Prasia Stal and Plautilla Stal (Boulard 1975),
however. In Prasta this character is restricted to
the males, but the literature does not record such
dimorphism for Plautilla).
(2) asharp ventrolateral fold in tergites 4-7 (more or
less distinct in all species) is shared by nearly all
species of Gymnotympana, but not found in rela-
ted genera.
(3) anenlarged distal part of male operculum. Both,
the broadly rounded shape (as found in B. bem-
melent, B. hamiltoni, and B. wegeneri), as the
triangular shape (as found in B. daviesi, B. for-
tuini, and B. pigrami), are found in
Gymnotympana, but do not occur in related ge-
nera.
(4) a very short, often rudimentary meracanthus
(found in all species with enlarged opercula) is
shared by Venustria Goding & Froggatt, 1904
and most species of Gymnotympana. This char-
acter might be coupled to the character descri-
bed immediately above.
(5) a laterally elongate basal part of male operculum
(in six species) is shared by nearly all Gymno-
tympana species and not found in other genera.
(6) abroad hyaline border along hind margin of teg-
men (in B. fortuini and B. loriae, fig. 84) is sha-
red by all species of Gymnotympana, Scottotym-
pana, and Venustria.
(7) an inwards curving ventral corner of tymbal,
with strongly inwards directed connecting bar
between abdomen and tymbal (only found in B.
fortuini) is found in most species of Gymno-
tympana, but not in other related genera.
(8) furthermore, all species except B. daviesi and B.
wegeneri have a more or less distinct colour pattern
of dark streaks and spots on head, pronotum, and
abdomen (figs. 8-9). This colour pattern is much
more distinct than in other species of Baeturia. An
even more distinct colour pattern is found in ma-
ny Gymnotympana species, in Scottotympana and
in two species of Chlorocysta Westwood, 1851.
Since these patterns are very variable between the
various species, it is not clear whether the patterns
found in the loriae group can be considered ho-
mologue with those in Gymnotympana.
The loriae group thus, seems to combine some ap-
parently convincing synapomorphies of Baeturia and
Gymnotympana. At this point of investigation, with
several groups of the ‘Baeturia and related genera
complex’ still undescribed, and with only vague indi-
cations as to the possible sister groups of the complex,
it is not possible to fully evaluate the meaning of the
character distribution in the /orize group. Current
phylogenetic analysis, involving all species of the
‘Baeturia and related genera complex’, however,
shows that a monophyletic loriae group as part of the
genus Baeturia, is the most parsimonious solution.
Furthermore, this solution is in accordance with the
distribution of all male genital characters.
Nevertheless, the remarkable number of conflic-
ting characters concentrated in the loriae group is
upsetting to any phylogenetic reconstruction.
Whatever generic allocation of the loriae group is fa-
voured, the parallel evolution and / or the successive
gain and loss of several characters must be presumed.
Biochemical study of the loriae group could be inte-
resting as a test for a hybrid origin of that group.
Monophyly of the B. loriae group
The above discussed combination of characters
found in the /oriae group involves all those characters
DE Boer: The loriae group of Baeturia
that would, by non-existence of that group, be used as
apomorphies for either Baeturia or Gymnotympana.
As a result neither Baeturia nor Gymnotympana can
be properly defined by shared apomorphies in exclu-
sion of the loriae group. But also for the loriae group
itself, no unambiguous apomorphies could be found.
The group can be best defined by the unique combi-
nation of two characters: sexual dimorphism in wing,
and an S-curved aedeagus with lateral lobes at the ba-
se of the curvation. Furthermore, there are three char-
acters shared by all, or nearly all, species of the group,
and not widely distributed outside the group, that
could indicate a close relationship between the spe-
cies. These characters are: 1) a distinct crest between
the lateral lobes of aedeagus, only shared by some spe-
cies of the B. conviva group and two species related to
that group; 2) an enlarged and angularly rounded
ventral part of the lateral lobe of pygofer, shared by
only a few species of Gymnotympana (e.g. G. stridens);
and 3) a rather short and broad eighth apical area of
tegmen, tending to be shorter than in Gymnotympana
or other species of Baeturia. The latter character is so-
mewhat variable within the species and cannot be
used as a diagnostic character.
Ingroup phylogeny
As long as the problems concerning the phylogene-
tic position of the loriae group, relative to other spe-
cies groups of Baeturia and to the genus
Gymnotympana, are not satisfactorily solved, charac-
ters that occur either in Baeturia or Gymnotympana
and also in some species of the loriae group cannot be
properly evaluated as plesiomorphous or apomorp-
hous, and can therefore not be used in a phylogenetic
reconstruction of the group. Only the distribution of
characters that appear to be unique for some species
of the loriae group allows some remarks on ingroup
phylogeny.
B. bemmeleni, B. hamiltoni, and B. wegeneri almost
certainly form a monophyletic group. These species
share an almost identical clasper, characterized by a
nearly rectangular dorsal corner (figs. 11, 26, and 30),
which is regarded synapomorphic. Furthermore,
these species share an almost identical male opercu-
lum; very broad, slightly domed and broadly rounded
at its apex (figs. 14, 22, and 38). A very similar oper-
culum was found in G. olivacea Distant, 1905, but in
that species the lateral margin of operculum is almost
continuous with the lateral crest of operculum base.
Some undescribed Gymnotympana species, related to
G. olivacea, have a similarly broad and curved, but
more angular, square-shaped, operculum.
B. daviesi, B. fortuini, B. hartonoi, B. loriae, B. pi-
grami, and B. silveri, most probably form a monophy-
letic group on account of a shared dorsal protrusion
on the clasper. This protrusion is partly, or complete-
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
136° 140°
144°
wbemmeleni 0°
A hamiltoni
@hartonoi
» etenuispina
Bwegeneri
[een]
148°
Fig. 1. Localities of Baeturia bemmeleni, B. hamiltoni, B. hartonoi, B. tenuispina, and B. wegeneri.
ly fused with the upwards curving proximal part of
the dorsal margin of clasper, and merges with the
clasper base, were in many other Baeturia species the
clasper heel is formed. B. tenuispina Blôte has a simi-
larly shaped protrusion at the corner of its clasper heel
(fig. 97) and should possibly be included in this
group.
In B. daviesi, B. pigrami, and B. silveri the dorsal
protrusion is very distinct and finger-shaped (figs. 40,
50, 64); in B. hartonoi, this protrusion is more angu-
lar and strongly curved mesiad (fig. 70); and in B. for
tuini it is rudimentary and only recognizable in the
specimens from Mt. Missim (fig. 79). The dorsal pro-
trusion is believed to be lost in the lectotype of B. lor
iae and the remaining specimens of B. fortuini.
The clasper of these six species is very broad in lat-
eral view; the dorsal margin of the clasper curves up-
wards from clasper base, so that no clasper heel is
formed. This clasper-shape is probably also synapo-
morphous.
B. fortuini, B. loriae, and B. hartonoi possibly form
a monophyletic group on account of the shared elon-
gate lateral lobes of aedeagus (figs. 73, 82, 92), and
the very broad, angular and laminiform clasper.
B. fortuini and B. loriae are very closely related and
probably sister species. They share triangular bronzed
spots in the tegmina (fig. 84) and a very long and
slender aedeagus (figs. 82, 92). Possibly these two
species are synonyms, and the lectotype of B. loriae is
just an aberration.
The relationships between B. daviesi, B. pigrami,
and B. silveri are less clear. These species share a trian-
gular clasper, with convex dorsal margin (lateral
view), which must probably be regarded more primi-
tive than the squarely laminiform clasper of the three
species mentioned above. Especially the claspers of B.
daviesi and B. pigrami are very similar, sharing a
broadly rounded and outcurving dorsal lobe, but the
clasper of B. hartonoi has a very similar outcurving
lobe. B. pigrami and B. silveri are conspicuous by the
brown patches along veins of tegmina. But B. guttuli-
nervis Blöte and B. guttulipennis Blöte, two otherwise
very different and not closely related species, have
similar patches.
Table 1. Altitudinal ranges of the species of the B. loriae
group in meters above sea level.
B. bemmeleni 35-1350
B. fortuini 1100-2100
B. hamiltoni 50-1200
B. hartonoi 100
B. loriae 1300
B. pigrami 120-1260
B. silveri 1800-2590 (8500 ft)
B. wegeneri 250
® daviesi
e fortuini
a loriae
v pigrami
A Silveri
Fig. 2. Localities of Baeturia daviesi, B. fortuini, B. loriae, B.
pigrami, and B. silveri.
BIOGEOGRAPHY
The B. loriae group is endemic to New Guinea,
and distributed in the northeastern part of Irian Jaya,
and in central and eastern Papua New Guinea. Many
specimens come from, or close by, the central and
eastern mountain ranges. The group seems to consist
mainly of montane species, probably with very res-
tricted areas of distribution, often reaching altitudes
of over 1000 m (table 1).
It is remarkable, that the subdivision made above,
between B. bemmeleni, B. hamiltoni, and B. wegeneri
versus the remaining species, is a subdivision between
a mainly western and a mainly eastern species group
(figs. 1, 2).
TAXONOMY
Description of the B. loriae group
The species of the B. loriae group are predominant-
ly brown coloured, sometimes greenish tinged. B. we-
generi and B. daviesi are densely speckled with small
brown spots, the other species are conspicuous by a
colour pattern of regularly distributed dark markings.
These species generally have two dark spots or longi-
tudinal streaks between eyes and lateral ocelli and of-
ten darkened lateral corners of postclypeus; a light
immaculate middorsal band on pronotum, someti-
DE Boer: The loriae group of Baeturia
mes bordered by almost black lines, and dark brown
streaks in and along the oblique fissures. The dark
markings on the mesonotum form two semi-circular
paramedian spots, at pronotal collar, two blackened
dents in front of cruciform elevation and lateral
bands, converging from pronotal collar to edges of
elevation. The cruciform elevation and a triangular
area in front of the elevation is generally light och-
raceous and unmarked. On the male abdomen, dark
markings tend to form dorsal, and sometimes lateral
bands, while a latero-ventral row of dark spots is often
very clear. The dorsal markings tend to be interrupted
middorsally by a narrow lighter coloured line. Ventral
side of male abdomen generally light ochraceous, seg-
mental hind margins often red. Female abdomen mo-
re irregularly dark spotted all over, though the light
middorsal band is often more conspicuous.
Females are shorter than males (on average), but
have a larger, more robust head and thorax and often
longer tegmina.
Head in dorsal view (fig. 3): Postclypeus distinctly
protruding beyond vertex lobes and broadly rounded
at frontal margin. Postclypeus 1.2-2.2 X as broad as
long and 0.5-0.7 X as broad as distance between eyes,
strongly varying within the species. Head 0.7-0.8 X
as wide as pronotal collar and only slightly narrower
than anterior width of pronotum. Vertex narrow,
ocelli large and close together. Distance between la-
teral ocelli less than 1.5 X the width of frontal ocellus
and 0.7-1.4 X as long as distance between eye and la-
teral ocellus. Eye 0.6-0.8 X as wide as distance be-
tween eyes. Head 0.9-1.2 X as long and 2.2-2.6 X as
wide as distance between eyes.
Legs: Ochraceous, sometimes with longitudinal
brown stains on femora and tibia. Fore femur (fig.
19) with row of three sharply pointed spines, dimi-
nishing in length towards tibia. Proximal spine about
as long as distance to middle spine. Middle spine
broad, triangular.
Tegmina and wings: Hyaline, though tegmina in
some species with brown markings. Venation och-
raceous, costa often reddish. Tegmina with 8, wings
with 6 apical areas. Costal area of tegmen very nar-
row. Anal fields of wing in male distinctly narrower
than in female (compare figs. 59 and 84).
Tymbal organ (fig. 6): Six or seven parallel trans-
verse sclerotized ridges spanning the tymbal from
dorsal to ventral margin. Short intercalary ridges for-
ming a midlateral band across tymbal.
Opercula: Basal part of operculum vaulted, with
distinct crest around rectangular distolateral corner.
Distal part of male operculum either very long and
domed, and completely covering tymbal cavity in
ventral view; or short, oblong-shaped, laminiform
and flat against body, only partly covering tymbal ca-
vity. Distal part of operculum medially extending
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 3-7. Baeturia hamiltoni sp. n. — 3, Head in dorsal view; 4, postclypeus in lateral view; 5, female genital segment in later-
al view; 6, tymbal; 7, female operculum.
DE Boer: The loriae group of Baeturia
Fig. 8. Baeturia hamiltoni sp. n., habitus male.
beyond meracanthus. Distal part of female opercu-
lum short, sickle-shaped and erect, medially exten-
ding just to base of meracanthus. Basal part of female
operculum forming narrow rim around base of mer-
acanthus.
Abdomen: With tergites 4-7 of the male abdomen
sharply folded at the ventrolateral edge of abdomen,
though these folds are not always distinct an all speci-
mens.
Male genitalia: Pygofer with stout and often angu-
larly bent caudodorsal beak. Lateral lobes of pygofer
with small and bluntly rounded protuberances, that
do not extend beyond pygofer margin. Lateral lobes
Fig. 9. Baeturia hamiltoni sp. n., habitus female.
strongly folded inwards. Claspers parallel and direc-
ted posteriad, slightly bending down towards their
apices. Apical part of clasper with small, rounded and
sharply edged clasper hollow. Aedeagus strongly S-
curved, with two narrowly elongate lateral lobes at ba-
se of S-curvation, and a distinct crest between lateral
lobes. Aedeagus in lateral view broad at base, gradual-
ly narrowing to pointed apex.
Female genitalia: Caudodorsal beak sharply poin-
ted at apex. Ovipositor sheats reaching just beyond
apex of beak (fig. 5).
Etymology: This revision is to form part of an area
cladistic study of the New Guinean region, in which
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
the general patterns of distribution of organisms are
compared to the geological history of that island. The
new species described in this publication are named
in honour of geologists, who contributed to a pale-
ogeographic reconstruction of the area. The species
are named after R. W. van Bemmelen, H. L. Davies,
A. R. Fortuin, W. B. Hamilton, H. M. S. Hartono,
C. J. Pigram, E. A. Silver, and A. Wegener.
Key to the males
1. Clasper with distinct angular clasper heel; apical
part of clasper slender in lateral view, with rectan-
gular dorsal corner (fig. 11). Distal part of oper-
culum broad and long, weakly domed, and bro-
adlyssoundedataperx (Hear m... 2
— Clasper without clasper heel; apical part of clasp-
er in lateral view broader than clasper base, often
with dorsal protrusion, partly fused to proximal
part of dorsal margin (fig. 40). Distal part of op-
erculum either oblong and flat (fig. 41), or trian-
gular, domed and pointed (fig. 51) at apex … … 3
2. Body densely speckled all over, not forming an
unspeckled or lighter coloured middorsal band ..
vic ti MeN eN B. wegeneri
— Body not speckled, but with regularly distributed
dark markings and an immaculate middorsal
band over part or whole length of body ............ 4
3. Body length 25.4-29.7 mm. Caudodorsal beak
broad and truncate at apex ............. B. hamiltoni
— Body length 17.1-21.3 mm. Caudodorsal beak
narrowly truncate or pointed at apex …
DIR POR Cee UD NE B. bemmeleni
4. Aedeagus very long, with elongate and slender
lateral lobes (fig. 82). Clasper quadrangular in
lateral view; dorsal part of clasper angular, lamin-
tormi(fes 707980889) en 5
— Aedeagus short and broad, lateral lobes not elon-
gate (fig. 45). Clasper triangular in lateral view;
dorsal part of clasper swollen, with weakly convex
dorsal margin (figs 40, 50, 64) … ii 7
5. Postclypeus not swollen in lateral view. Tegmen
immaculate. Aedeagus angularly bent to apex
(fig. 73). Clasper with dorso-lateral lobe (fig. 70)
B. hartonoi
— Postclypeus distinctly swollen in lateral view.
Tegmen with triangular brown spots in apical ar-
eas (fig. 84). Aedeagus not angularly bent to apex
(fig. 81). Clasper without dorso-lateral lobe (fig.
TD) ER SI MEINEN ER 6
6. Distal part of operculum triangular, domed and
narrowly rounded at apex (fig. 77). Margin of ae-
deagus pore concave at apex (fig. 83) .B. fortuini
— Distal part of operculum square-shaped, flat
against body (fig. 91). Margin of aedeagus pore
not concave at apex (fig. 93) .................. B. loriae
7. Tegmina with brown markings along veins (fig.
DOME ER NN e ese ee 8
=) egminaimmaculateweres een B. daviesi
8. Postclypeus angularly swollen in lateral view.
Distal part of operculum triangular and slightly
domed. Hind margin of tegmen with distinct
hyaline borde ee B. pigrami
— Postclypeus not swollen in lateral view. Distal
part of operculum square-shaped and flat against
body. Hind margin of tegmen with narrow hya-
line border eee ER ne B. silver:
Description of the species
Baeturia hamiltoni sp. n.
(figs. 1, 3-20)
Type material. — Holotype d: ‘Neth. Ind.-American
New Guinea Exped. Rattan camp, 1200 m, 6.iii.1939, L. J.
Toxopeus’, RMNH. - Paratypes: IRIAN JAYA: NEW GUINEA
(W): Araucaria camp, 800 m, 8.iii.1939, L. J. Toxopeus,
16, 39, RMNH; same data but 9.iii.1939, 16, 19;
10.11.1939, 32; 11.11.1939, 16; 12.11.1939, 19;
16.11.1939, 16, 29; 20.11.1939, 19; 21.11.1939, 26;
22.11.1939, 26, 29; 24.11.1939, 29; 25.11.1939, 18;
29.11.1939, 16, all RMNH; same data but 19.11.1939, 29;
28.11.1939, 28, all ZMA; Bernhard camp, 100 m,
11.iv.1939, L. J. Toxopeus, 22, RMNH.
Other material. — PAPUA: NEW GUINEA (NE): Ambunti,
Sepik R., 50 m, 10.v.1963, R. Straatman, 1d, BPBM.
B. hamiltoni is a large brown coloured, species with a light
middorsal band on head and thorax, sometimes continuing
over abdomen. The species is easily recognized by a small
and almost black medial spot on pronotum, at pronotal col-
lar. Females have a glistering middorsal band of silvery setae
on the abdomen, making the light and immaculate middor-
sal band more conspicuous than in males.
Description
Body of males brown, abdomen sometimes red-
dish, with distinct colour pattern of darkened mar-
kings and an immaculate middorsal band on head
and thorax, sometimes continuing over abdomen.
Females on average slightly smaller than males, with
more robust head and thorax, but shorter abdomen
and considerably longer tegmina. Male abdomen 1.3-
1.4X as long as head and thorax, of female 0.9-1.0 x.
Tegmina of males 1.1-1.2 as long as total body
length, of females 1.4-1.5 x.
Head (fig. 3): Light brown, with traces of red
around ocelli, longitudinal dark brown to black
streaks on vertex lobes, between eyes and lateral ocel-
li, and often brown spots at margins of postclypeus.
Postclypeus very stout, 1.3-1.9 as long as broad,
distinctly protruding beyond vertex lobes and broad-
ly rounded at anterior margin. Postclypeus in lateral
view (fig. 4) strongly inflated, and angularly protrud-
ing; dorsal part of anterior margin (lateral view) al-
DE Boer: The loriae group of Baeturia
20
Figs. 10-20. Baeturia hamiltoni sp. n. — 10, pygofer in lateral view; 11, clasper in lateral view; 12, clasper in dorsal view; 13,
male caudodorsal beak in dorsal view; 14, male operculum; 15, aedeagus from aslant; 16, aedeagus in lateral view; 17, aedea-
gus from behind; 18, detail crest between lateral lobes of aedeagus; 19, fore femur; 20, female caudodorsal beak in dorsal view.
Lettering: bp = basal part of operculum; c = crest between lateral lobes of aedeagus; cb = caudodorsal beak; dm = distomedi-
al margin of operculum; di = distal margin of pygofer; dc = crest around distolateral corner of basal part of operculum; do =
dorsal margin of pygofer; dp = distal part of operculum; he = clasper heel; ho = clasper hollow; la = lateral margin of opercu-
lum; m = meracanthus; me = medial margin of operculum; p = protuberance on lateral lobe of pygofer; ve = ventral margin
of pygofer.
most straight, ventral part strongly concave. Head
very narrow between eyes, distance between lateral
ocelli only slightly larger than width of frontal ocel-
lus. Distance between lateral ocelli 0.8-1.2 x distance
between eye and lateral ocellus. Head narrower than
anterior part of pronotum.
Thorax: Pronotum brown, with broad dark brown
streaks, in and along oblique fissures, and broad, im-
maculate medial band. Markings variable in intensity
and shape, and sometimes completely absent. All
specimens with a small dark brown or black middor-
sal spot at margin of pronotal collar. Mesonotum
light brown, with two paramedian dark spots at pro-
notum margin, darkened lateral streaks converging
from pronotal collar to corners of cruciform elevation
and two black spots in front of elevation. Cruciform
elevation, and a narrow triangular area in front of el-
evation, often reaching to pronotal collar, light och-
raceous and immaculate.
Legs: Fore femur (fig.19) with row of three erect
spines, diminishing in length towards tibiae.
Tegmina and wings: Hyaline, though slightly
bronzed, venation ochraceous or reddish tinged, but
costa often bright red. Veins densely set with short se-
tae. Tegmina with very short apical areas, 8th area al-
most square-shaped, and with a very narrow hyaline
border along hind margin. This border slightly broad-
er in wings.
Tymbal organs (fig. 6): Tympanum not curving in-
wards; connecting bar between tymbal and abdomen
quite short and almost parallel to body axis. Six trans-
verse sclerotized ridges spanning the tymbal, from
dorsal to ventral margin. A 7th ridge, close to proxi-
mal tymbal margin, almost reaching ventral margin.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Six short intercalary ridges seem to form a band across
tymbal.
Opercula: Male operculum (fig. 14) ochraceous
brown with darkened streaks. Basal part of opercu-
lum almost oblong-shaped, but at half-width slightly
longer than at lateral or medial margin. Distal part of
operculum very broad, shell-shaped, and almost flat
against body. Operculum broadly rounded at apex
and reaching to, or beyond, margin of abdominal seg-
ment 3. Distomedial margin and distal % of lateral
margin convex, proximal % of lateral margin straight.
Medial margin short and straight, recurving to base of
meracanthus. Opercula well separated medially; 1st
sternite slightly elevated between opercula, with dis-
tinct distal lobe. Meracanthus very short, reaching to
about % the operculum length. Female operculum
(fig. 7) sickle-shaped and erect, with weak crest along
distal margin; area between operculum and base of
meracanthus broad and flat.
Abdomen: Male abdomen light brown, with some
darker brown spots on lateral sides and latero-ventral
rows of dark spots on segments 3-7. Segments 2-7
darkened dorsally as well, though these markings of-
ten interrupted by a narrow lighter middorsal band.
Segment 8 almost entirely dark brown, but pygofer
ochraceous. Ventral side of abdomen light brown,
immaculate. Segmental hind margins often reddish.
Anterior margin of 2nd tergite convex medially; ante-
rolateral sides of 2nd tergite distinctly swollen adja-
cent to tymbal. Female abdomen more irregularly
spotted all over. Light middorsal band more conspic-
uous than in males, often glistering with short, silvery
setae. Ovipositor sheaths reaching just beyond apex
of caudodorsal beak (fig. 5). Female caudodorsal beak
(fig. 20) stout, slightly erect and rounded at apex.
Male genitalia: Pygofer slender in lateral view as in
fig. 10. Dorsal margin straight, angularly bending
into stout, short and slightly erect caudodorsal beak.
Distal margin straight angularly bending into margin
of beak. Lateral lobes of pygofer strongly curved in-
wards, with bluntly rounded lateral protuberances.
Ventral margin angularly convex, but concave near
base of pygofer. Caudodorsal beak in dorsal view (fig.
13) broad and truncate at apex. Clasper in lateral view
(fig. 11) broad at base, with very distinct and angular
clasper heel, strongly bent down at half-length, form-
ing almost right angled dorsal corner. Dorsal margin
of clasper concave between clasper heel and this dor-
sal corner. Distinct crest running along % of dorsal
margin, and bending outwards around aedeagus to-
wards clasper base. Clasper hollow short, broadly
rounded at apex. Clasper in dorsal view (fig. 12), with
dorsal margins of clasper bases very close together,
leaving only a narrow gap for anal valves. Aedeagus in
lateral view (fig. 16) strongly concave around aedea-
gus pore, lateral lobes long and slender. Distinct crest,
10
at base of S-curvation, between lateral lobes (figs. 15,
18). Aedeagus pore (fig. 16) large and oval-shaped.
Measurements: Body length ó: 25.4-29.7 mm (x
27.6 mm + 1.3), 2: 23.5-28.7 mm (x 26.6 mm +
1.4); tegmen length d : 28.0-35.0 mm (x 32.5 mm +
1.8), 2: 32.1-40.5 mm (x 37.7 mm + 2.3); head
length d: 2.3-2.6 mm (x 2.4 mm), 9: 2.4-3.1 mm (x
2.6 mm); pronotum length d: 3.1-3.8 mm (x 3.5
mm), 2: 3.5-4.3 mm (x 4.0 mm); mesonotum length
d: 5.8-6.6 mm (x 6.3 mm), 9: 6.5-7.8 mm (x 7.1
mm); head width d: 5.5-6.2 mm (x 5.8 mm), 9:
5.9-6.9 mm (x 6.6 mm); width of pronotal collar
36: 7.1-8.5 mm (x 8.0 mm), 9: 8.3-9.9 mm (x 9.2
mm).
Distribution (fig. 1). — B. hamiltoni is known from
a very restricted area in Irian Jaya and from one local-
ity in NW Papua.
Etymology. — The species is named in honour of
the geologist W. B. Hamilton.
Remark. — Three females from Araucaria camp that
have been described as B. guttulinervis Blote (allo- and
paratypes) are very similar in size and colour pattern
to the females of B. hamiltoni, but have darkened
patches along the venation of tegmen, are slightly
brown speckled on the pronotum and miss the dark
middorsal spot at pronotal collar.
Baeturia bemmeleni sp. n.
(figs. 1, 21-29)
Type material. — Holotype d: ‘NEW GUINEA NETH.
Bodem, 100 m, 11 km SE of Oerberfaren, July 7- 17. 1959’;
‘T. C. Maa Collector’, BPBM. — Paratypes: IRIAN: NEW GUIN-
EA (W): Boven Digul gebied [Upper Digul area], 400 km N.
Merauke, Dr A. Kalthofen, legit 1926, 16, ZMA; Boven
Sermowai riv., + 400 m, 8.iv.1911, K. G. [K. Gjellerup],
26, RMNH.
Other material. — PAPUA: NEW GUINEA (NE): Eliptamin
valley, 1200-1500 m, 16-31.vii.1959, W.W. Brandt, 1 9,
BPBM; NEW GUINEA (SE): Kiunga, Fly riv., 35 m, viii.1969, J.
and M. Sedlacek, 1 9, BPBM.
B. bemmeleni closely resembles B. hamiltoni in
shape of male operculum and clasper, but is consider-
ably smaller and less intensely coloured. The female
from Eliptamin Valley deviates somewhat in shape
and colour pattern, but the shape of its 8th apical ar-
ea of tegmen suggests that this specimen belongs to
the B. loriae group and it presumably belongs to this
species.
Description
Body of males ochraceous or reddish brown, with
light brown patches, smaller than in foregoing spe-
cies. Distribution of patches resemble markings in B.
hamiltoni; leaving a light middorsal band on head and
DE Boer: The loriae group of Baeturia
ZN
24 | \ 25 26
30
Figs. 21-30. 21-29: Baeturia bemmeleni sp. n. — 21, pygofer in lateral view; 22, male operculum; 23, female operculum; 24,
male caudodorsal beak in dorsal view; 25, female caudodorsal beak in dorsal view; 26, clasper; 27, aedeagus from behind; 28,
aedeagus in lateral view; 29, detail crest between lateral lobes of aedeagus. — 30: Baeturia wegeneri sp. n., clasper.
thorax immaculate. This band sometimes interrupted
by a vaguely darkened middorsal spot, at margin of
pronotal collar. Females greenish brown, specimen
from Eliptamin Valley densely brown speckled. Both
females show the light brown middorsal marking on
pronotum. Male abdomen 1.3-1.6X as long as head
and thorax, of females 1.1 x. Male tegmen 1.1-1.2 x
as long as total body length, of females 1.3 x.
Head: Ochraceous, with brown markings on vertex
lobes and along sides of postclypeus, but densely
speckled in female of Eliptamin Valley. Postclypeus
strongly protruding, almost triangular in dorsal view,
1.8-2.3 x as wide as long. Postclypeus in lateral view,
angularly inflated as in B. hamiltoni, though in some
specimens more rounded at anterior margin. Head
about as wide as anterior part of pronotum. Distance
between lateral ocelli 0.7-0.9 as long as distance
between eye and lateral ocellus.
Thorax: Pronotum light brown, with dark brown
spots along oblique fissures and a light, but narrow,
middorsal band. Female from Eliptamin Valley
brown speckled between medial fissures and showing
this band only on proximal half of pronotum. Slightly
darkened middorsal spot, at margin of pronotal col-
lar, marking end of immaculate band; this spot less
clear than in B. hamiltoni. Spot U-shaped in one male
from Sermowai riv. and the female from Kiunga riv.;
connecting two dark lines, running partly along the
immaculate middorsal band. Mesonotum variable in
colour pattern between individual specimens, though
always with a light, immaculate cruciform elevation
and, with exception of the densely speckled female
from Eliptamin Valley, with a light middorsal band
or narrow triangular area in front of elevation. Two
dark spots in front of cruciform elevation.
Tegmina and wings: Hyaline, veins slightly red-
dened. Tegmina with short 8th apical area, and with
narrow hyaline border along hind margin. Wings
with slightly broader hyaline border.
Tymbal organs: As in B. hamiltoni, not curving in-
wards, but with connecting bar between tymbal and
abdomen distinctly longer and more curved inwards.
Six transverse sclerotized ridges spanning the tymbal,
from dorsal to ventral margin. A 7th ridge, close to
proximal tymbal margin, almost reaching ventral
margin. Six short intercalary ridges seem to form a
band across tymbal.
Opercula: Male operculum (fig. 22) closely resem-
bling that of B. hamiltoni, broadly rounded, shell-
shaped, and ochraceous brown with some darkened
streaks. Basal part of operculum almost oblong, but
weakly tapering mesiad at half its width. Distal part
weakly domed, almost flat against body, its rounded
apex reaching well beyond margin of abdominal seg-
ment 3. Distomedial margin and distal % of lateral
margin weakly convex, proximal % of lateral margin
more strongly convex. Meracanthus very short, reach-
ing to about % the operculum length. Female opercu-
wl
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 31-38. Baeturia wegeneri sp. n. — 31, pygofer in lateral view; 32, pygofer from aslant; 33, first sternite; 34, caudodorsal
beak in dorsal view; 35, aedeagus from behind; 36, aedeagus in lateral view; 37, detail crest between lateral lobes of aedeagus;
38, operculum.
lum (fig. 23) sickle-shaped and erect, with broad and
flat area between base of meracanthus and medial
edge of operculum.
Abdomen: Male abdomen ochraceous brown,
stained with dark brown spots, concentrated in near-
ly continuous middorsal band. Sternites immaculate.
A latero-ventral row of clear dark spots on segments
3-7. Segmental hind margins bright red. Anterior
margin of 2nd tergite convex medially; sides of 2nd
tergite distinctly swollen and adjacent to tymbal.
Female abdomen darker brown with irregular brown
markings. Latero-ventral row of dark spots less con-
spicuous than in males. Segmental hind margins och-
raceous. Female caudodorsal beak (fig. 25) long, slen-
der, and pointed at apex.
Male genitalia: Pygofer (fig. 21) closely resembling
that of B. hamiltoni, slender in lateral view. Dorsal
margin straight, angularly bending into straight, stout
and slightly erect caudodorsal beak. Distal margin
straight, angularly bending into margin of beak.
Lateral lobes of pygofer strongly incurved, with well-
developed, bluntly rounded lateral protuberances.
Ventral margin angularly convex, but concave near
base of pygofer. Caudodorsal beak (fig. 24) short,
sharply pointed or narrowly truncate at apex. Clasper
12
in lateral view (fig. 26) almost identical to that of B.
hamiltoni, with very distinct angular clasper heel, and
rectangular dorsal corner. Dorsal margin of clasper
concave between clasper heel and this dorsal corner.
Distinct crest running along % of dorsal margin, and
bending outwards around aedeagus towards clasper
base. Clasper hollow short, broadly rounded at apex.
In dorsal view, dorsal margins of clasper bases wider
apart than in B. hamiltoni. Aedeagus in lateral view
(fig. 28) slender, strongly concave along margin of
pore. Lateral lobes long and slender. Distinct crest at
base of S-curvation, between lateral lobes (fig. 29).
Aedeagus seen from behind (fig. 27) very slender.
Aedeagus pore narrow and oval.
Measurements: Body length d: 17.1-21.3 mm (x
18.4mmt+ 1.7), 2: 19.3 and 21.7 mm; tegmen length
3: 19.8-20.4 mm, 9: 27.8 and 28.6 mm; head length
3: 1.5-1.8 mm (x 1.7 mm), 2: 1.9 and 2.2 mm; pro-
notum length d: 1.9-2.2 mm (x 2.1 mm), ?: 2.8 and
3.2 mm; mesonotum length d: 3.4-3.8 mm (x 3.7
mm), 2: 5.0 and 5.3 mm; head width & : 4.2-4.5 mm
(x 4.3 mm), 2: 5.0 and 6.0 mm; width of pronotal
collar &: 5.2-5.4 mm (x 5.3 mm), ®: 6.9 and 7.2 mm.
Distribution (fig. 1). — B. bemmeleni is known
from several localities in the northern and central
parts of Irian Jaya.
Etymology. — The species is named in honour of
the geologist R. W. van Bemmelen.
Baeturia wegeneri sp. n.
(figs. 1, 30-38)
Type material. — Holotype d : ‘NEW GUINEA: NE, May R.
Petrol sta., 250 m, 3.vi.1963’ [print]; ‘Dry Forest’ [print];
‘R. Straatman Collector BISHOP’ [print], BPBM. — Paratype:
same locality and collector as holotype but 100 m,
31.v.1963, 16, BPBM.
B. wegeneri is an ochraceous brown coloured spe-
cies, with densely brown speckling all over its body, as
characteristic for many Baeturia species, and in this
respect quite different from most other species of the
B. loriae group. However, B. wegeneri is very similar
to the two foregoing species in shape of male opercu-
lum and genitalia.
Description
Body ochraceous brown, densely brown speckled
all over. Abdomen strongly inflated, 1.7 X as long as
head and thorax. Tegmina about as long as body
length.
Head: Ochraceous, brown speckled. Postclypeus
broad and oblong in dorsal view, 1.4-1.6X as broad
as long. Postclypeus angularly inflated ventrally (as in
fig. 4). Head narrower than anterior part of prono-
tum. Distance between lateral ocelli 0.8-1.0X dis-
tance between eye and lateral ocellus.
Thorax: Pronotum ochraceous, brown speckled in
middorsal band and on pronotal collar. Mesonotum
grey-brown, densely brown speckled. Two dark spots
in front of speckled cruciform elevation, almost lost
in speckling.
Tegmina and wings: Hyaline, veins ochraceous or
slightly reddish and set with short setae. Tegmen with
short and squarish 8th apical area, and narrow hyaline
border along hind margin. Wing with slightly broad-
er hyaline border.
Tymbal organs: Not curving inwards, connecting
bar between tymbal and abdomen quite short and al-
most parallel to body axis. Six transverse sclerotized
ridges spanning the tymbal, from dorsal to ventral
margin. A 7th ridge, close to proximal tymbal mar-
gin, almost reaching ventral margin. Six short interca-
lary ridges seem to form a band across tymbal.
Operculum (fig. 38): Broad and rounded as in
foregoing species, ochraceous with some darkened
stains near lateral margin. Basal part of operculum al-
most oblong, but weakly tapering towards mesiad at
half its width. Distal part slightly domed, almost flat
against body, broadly rounded at apex and reaching
DE Boer: The loriae group of Baeturia
just beyond margin of abdominal segment 3. Distome-
dial margin and distal # of lateral margin convex, prox-
imal % of lateral margin more strongly convex.
Opercula well separated medially, by distinctly smaller
and anteriorly lobate first sternite (fig. 33). Meracan-
thus short, reaching to about % of operculum length.
Abdomen: Ochraceous brown, tergites densely
brown speckled all over, sternites unspeckled. Latero-
ventral row of dark spots almost inconspicuous, by
speckling. Segmental hind margins bright red.
Anterior margin of 2nd tergite convex middorsally;
sides of 2nd tergite weakly swollen and adjacent to
tymbal.
Genitalia: Pygofer in lateral view (fig. 31) resem-
bling that of foregoing species. Dorsal margin straight,
angularly bending into stout and slightly erect caudo-
dorsal beak. Distal margin straight and more gradual-
ly bending into margin of beak. Lateral lobes of pygof-
er strongly curved inwards, with well-developed,
bluntly rounded, lateral protuberances. Ventral mar-
gin straight, not concave to base; ventral margins con-
verge to sharp angle at base of pygofer opening (fig.
32). Caudodorsal beak slightly longer than in B. bem-
meleni, very slender in dorsal view (fig. 34) and sharp-
ly pointed apically. Clasper in lateral view (fig. 30)
strongly resembling that of two foregoing species, with
a very distinct angular clasper heel and rectangular
dorsal corner. Clasper strongly bent down to apex at
half-length. Distinct crest running along % of dorsal
margin. Clasper hollow short and broadly rounded at
apex. Dorsal margins of clasper bases wider apart than
in B. hamiltoni. Aedeagus in lateral view (fig. 36) con-
cave along margin of pore, with broader basal lobes
than in B. bemmeleni. Distinct crest at base of S-curva-
tion, between basal lobes (fig. 37). Aedeagus from be-
hind (fig. 35) broader than in B. bemmeleni, with a
short and broad oval pore.
Measurements: Body length: 26.0 and 27.3 mm;
tegmen length: 26.4 and 26.6 mm; head length: 2.0
and 2.3 mm; pronotum length: 2.6 and 2.7 mm;
mesonotum length: 5.3 mm; head width: 4.7 and 4.8
mm; width of pronotal collar: 6.4 and 6.5 mm.
Distribution (fig. 1). — B. wegeneri is only known
from May river, in central west Papua New Guinea.
Etymology.— The species is named in honour of
the geologist A. Wegener.
Baeturia silveri sp. n.
(figs. 2, 39-48)
Type material. — Holotype d: ‘NEW GUINEA: NE Purosa,
20-26 km SE Okapa, 1800-2020 m, 28.viii.1964 [print];
‘J. and M. Sedlacek Collectors BISHOP’ [print], BPBM. —
Paratypes: same data as holotype 16, 12, BPBM.
Other material. — PAPUA: NEW GUINEA (NE): Kandep,
13
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
42
43
48
Figs. 39-48. Baeturia silveri sp. n. — 39, pygofer in lateral view; 40, clasper; 41, male operculum, Kandep; 42, male caudodor-
sal beak in dorsal view; 43, female caudodorsal beak in dorsal view; 44, aedeagus from behind; 45, aedeagus in lateral view;
46, detail crest between lateral lobes of aedeagus; 47, male operculum, holotype; 48, female operculum. Lettering: d = distal
margin of operculum; | = lateral margin of operculum; m = medial margin of operculum.
Western Highlands, 8000 ft, 23.xii.1961-14.ii.1962, W. W.
Brandt, 1d, CSIRO.
B. silveri can be recognized by the brown speckled
tegmina and short, oblong male operculum.
Description
Body of males light brown or ochraceous, with a
colour pattern of dark brown spots and a narrow, im-
maculate, middorsal band as characteristic for most
species of the B. loriae group. Female slightly reddish
brown with irregularly distributed dark spots. Male
abdomen 1.4-1.6 as long as head and thorax, of fe-
male 1.1 x. Tegmina of males 1.2-1.4 x as long as to-
tal body length, of female 1.5.
Head: Ochraceous brown, with dark brown streaks
on vertex lobes, almost black between eyes and lateral
ocelli. Female with black markings between prono-
tum and ocelli. Postclypeus sometimes darkened to-
wards lateral margins, 1.6-1.9 x as broad as long and
broadly rounded at anterior margin. Postclypeus not,
or only slightly, swollen ventrally, anterior margin
(lateral view) straight or weakly convex. Head nar-
rower than anterior part of pronotum. Distance
between lateral ocelli 0.9-1.3 x distance between eye
and lateral ocellus.
14
Thorax: Pronotum light brown, darkened along
fissures; with a lighter, ochraceous, middorsal band,
sharply bordered by irregular black streaks. Middorsal
band abruptly widening near pronotal collar, and
ending at a vaguely darkened middorsal spot.
Middorsal band reddish brown and brown speckled
in the female, but clearly recognizable by its black lin-
ing. Pronotal collar red-brown, brown speckled.
Mesonotum with two distinct. semi-circular dark
spots at pronotum margin and darkened, greenish
tinged, broad lateral streaks, converging from prono-
tum margin to corners of cruciform elevation.
Elevation and narrowly pointed triangular area in
front of elevation, light ochraceous. Mesonotum of
female, including cruciform elevation, reddish tinged
and densely brown speckled.
Tegmina and wings: Hyaline, venation reddish
ochraceous. Tegmen with irregular brown markings,
predominantly along venation. Hyaline border along
hind margin of tegmen broader than in foregoing
species and almost as broad as in wing. The 8th apical
area of tegmen slightly longer, more slender, than in
other species of this group.
Tymbal organ: Not curving inwards, connecting
bar between tymbal and abdomen quite short and al-
most parallel to body axis. Five transverse sclerotized
ridges spanning the tymbal, from dorsal to ventral
margin. A 6th ridge almost reaching ventral margin,
and a 7th, most proximal, ridge running from dorsal
margin to about half the tymbal width. Six intercalary
ridges seem to form a band across tymbal. Traces of
red between tymbal ridges.
Opercula: Male operculum (fig. 47) quite different
from that of the three foregoing species, more resem-
bling the opercula found in other groups of Baeturia.
Basal part of operculum oblong, equally long at later-
al and medial margins. Distal part angularly oblong,
wider than long and flat against the body, almost
completely covering tymbal cavity. Lateral and medi-
al margin almost straight, distal margin weakly con-
vex. Distolateral corner broadly rounded, distomedial
corner angular. Opercula wide apart medially, separ-
ated by broad and rounded Ist sternite. Meracantus
almost as long as operculum. Operculum of Kandep
specimen (fig.41) much longer and more directed
mesiad, with longer lateral margin. Female opercu-
lum (fig. 48) sickle-shaped and erect. Distal margin
convex.
Abdomen: Male abdomen light brown, ochraceous
in Kandep specimen. Two parallel dorsal rows of dark
spots along a narrow light middorsal band and a very
clear latero-ventral row of dark spots on segments 3-
7. Lateral and ventral sides of abdomen slightly
brown stained. Abdomen of Kandep specimen with
midlateral band of dark spots, as in B. fortuini and B.
loriae. Anterior margin of 2nd tergite convex medial-
ly; sides of 2nd tergite distinctly swollen and adjacent
to tymbal. Female abdomen irregularly brown spot-
ted all over, though with unstained light middorsal
band. Female caudodorsal beak (fig. 43) pointed at
apex.
Male genitalia: Pygofer in lateral view rounded,
with stout caudodorsal beak (fig. 39). Dorsal margin
straight almost continuous with erect caudodorsal
beak. Distal margin straight or slightly concave,
broadly rounded into margin of beak. Lateral lobes of
pygofer curved inwards, with well-developed, bluntly
rounded protuberances, that curve slightly upwards
towards pygofer margin. Ventral margin angularly
convex, but concave near base of pygofer.
Caudodorsal beak (fig. 42) rounded, almost truncate
at apex. Clasper in lateral view (fig. 40) strongly
rounded and quite different from the three foregoing
species. Clasper gradually widening distad to clasper
base, so that no clasper heel is formed. Dorsal margin
of clasper rounded, forming an almost globular,
slightly inwards curved, protrusion at about % its
length and, distad to this protrusion, convex to apex.
Clasper slightly curved outwards, towards dorsal mar-
gin and around aedeagus. Clasper hollow narrow tri-
angular, narrowly rounded at apex. Aedeagus longer
DE Boer: The loriae group of Baeturia
than in foregoing species, in lateral view (fig. 45)
strongly concave along margin of pore, with long and
slender basal lobes. Distinct crest at base of S-curva-
tion between basal lobes (fig. 46). Aedeagus pore (fig.
44) large and oval-shaped, but truncate at apex.
Measurements: Body length d: 20.0-22.0 mm (x
20.9 mm), 9: 18.8 mm; tegmen length d : 25.9-28.8
mm (x 27.2 mm), ®: 28.6 mm; head length d: 1.5-
1.7 mm, 9: 1.8 mm; pronotum length d: 2.2-2.5
mm, ®: 2.5 mm; mesonotum length d 4.2-4.6 mm,
2: 5.0 mm; head width 6: 3.8-4.3 mm, 2: 4.4 mm;
width of pronotal collar d : 5.4-6.2 mm, 9: 6.2 mm.
Distribution (fig. 2). — B. silveri is known from on-
ly two localities, both in the central mountain ranges
of Papua New Guinea.
Etymology. — The species is named in honour of
the geologist E.A. Silver.
Baeturia pigrami sp. n.
(figs. 2, 49-59)
Type material. — Holotype d: ‘Museum Leiden Neth
New Guinea Exp. Star Range, 1260 m, Sibil, 21.viii.1959,
op licht’ [print], RMNH.
Other material. — PAPUA: NEW GUINEA (NE): Feramin,
150-120 m, 1-6.vi.1959, W.W. Brandt, 22, BPBM.
Of this species only one male is available, it resem-
bles B. silveri in colour patterns of body and tegmina,
and in clasper shape. However, B. pigrami is much
larger, has very large triangular opercula, and an an-
gularly swollen postclypeus. Two females, with simi-
lar colour patterns and from the same area, probably
belong to this species.
Description
Body of male dark reddish brown, with very dis-
tinct light middorsal band on head and thorax. This
band much broader in females, and continuing over
abdomen. Male abdomen strongly inflated and 1.5 x
as long as head and thorax of females 1.0 X. Tegmina
of male 1.2 x as long as total body length, of females
1.4-1.5 X.
Head: Light brown with dark stains on vertex lobes,
mainly between lateral ocelli and eyes. Vertex and post-
clypeus with long setae, more densely set than in fore-
going speces. Postclypeus unstained, light ochraceous
in females, slightly brown suffused in male. Male post-
clypeus 1.2X as broad as long, in females 1.5 and
2.2 x. Postclypeus angularly swollen, almost conical in
lateral view (as in fig. 4). Vertex in male very narrow,
with ocelli close together. Head narrower than anterior
part of pronotum. Male head 1.2 x as long as distance
between the eyes, in female 0.9. Distance between
lateral ocelli in male 0.7 X distance between eye and
lateral ocellus, in females 1.1-1.2 x.
19
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 49-58. Baeturia pigrami sp. n. — 49, pygofer in lateral view; 50, clasper; 51, male operculum; 52, aedeagus from behind;
53, aedeagus in lateral view; 54, detail crest between lateral lobes of aedeagus; 55, male caudodorsal beak in dorsal view; 56,
female operculum; 57, female caudodorsal beak in dorsal view; 58, tymbal.
Thorax: Pronotum dark brown with light ochra-
ceous middorsal band, narrow and sharply defined in
male, much broader and more diffuse in females.
Pronotal collar slightly reddish and, especially in
male, stained with brown. Male mesonotum dark
brown, but light ochraceous in triangle-shaped area
in front of reddish brown cruciform elevation.
Female mesonotum with continuous broad ochra-
ceous middorsal band. Two black spots in front of
cruciform elevation in male and female.
Tegmina and wings (fig. 59): Hyaline, though teg-
mina with irregular brown markings, predominantly
along veins, venation reddish brown. Eight apical ar-
ea short and broad, especially in females. Tegmen and
wing with very narrow hyaline border along hind
margin.
Tymbal (fig. 58): Five transverse sclerotized ridges
spanning the tymbal from dorsal to ventral tymbal
margin. A 6th ridge almost reaching ventral tymbal
margin, and a 7th, most proximal, ridge running
from dorsal margin to only about half the tymbal
width. Six intercalary ridges seem to form a band
16
across tymbal. Traces of red between tymbal ridges.
Opercula: Male operculum (fig. 51) very large, red-
dish ochraceous with dark stains. Basal part of oper-
culum with greatest length at % its width, slightly ta-
pering towards lateal margin, more strongly tapering
to base of meracanthus. Distal part triangle-shaped
and slightly domed, narrowing towards narrowly
rounded, almost pointed apex and reaching to half-
length 3rd abdominal segment. Lateral margin of
male operculum convex near base, concave at half-
length, and weakly convex towards apex. Distomedial
margin concave near apex and convex to short and
straight medial margin. Opercula close together me-
dially, separated by narrow and weakly elevated 1st
sternite. Meracanthus very short, reaching to about %
the operculum length. Female operculum (fig. 56)
sickle-shaped, slightly angular at distal margin, and
erect. Base of operculum forming narrow rim around
base of meracanthus.
Abdomen: Male abdomen brown, dark brown
stained dorsally and on sides of 7th and 8th segments,
though lighter and reddish tinged, in very narrow
DE BOER: The loriae group of Baeturia
Fig. 59. Baeturia pigrami n. sp., female tegmen and wing.
middorsal band. A latero-ventral row of slightly dark-
ened patches, hardly conspicuous. Segmental hind
margins slightly red. Anterior margin of 2nd tergite
convex middorsally. Lateral parts of 2nd tergite, with
auditory capsules, and anterolateral part of 3rd ter-
gite, weakly incurving towards ventral corner of tym-
bal. Tergites 4-7 with sharp latero-ventral fold.
Female abdomen ochraceous, densely stained with ir-
regular brown spots laterally and ventrally, but with
broad and immaculate middorsal band. Female cau-
dodorsal beak very characteristic in dorsal view (fig.
57), with greatest width at about % its length and
sharply pointed at apex.
Male genitalia: Pygofer in lateral view (fig. 49).
Dorsal margin of pygofer bending gradually into erect
caudodorsal beak. Distal margin straight, angularly
bending into margin of beak. Lateral lobes of pygofer
strongly curving inwards, with well-developed, blunt-
ly rounded protuberances. Ventral margin angularly
convex, but strongly concave to base of pygofer.
Caudodorsal beak in dorsal view (fig. 55) very slender,
narrowly rounded, almost pointed at apex. Clasper
(fig. 50) closely resembling that of B. silveri, not
forming a clasper heel and slightly bent down to apex.
Dorsal margin of clasper strongly bent upwards from
clasper base, forming a long finger-shaped, slightly in-
curved protuberance. This protuberance distinctly
longer than in B. silveri. Clasper strongly bent out-
wards distally of this protuberance, and forming a
broadly rounded dorsal crest. Clasper hollow longer
than in B. silveri. Aedeagus in lateral view (fig. 53)
slightly concave along margin of pore, with narrow
and elongate lateral lobes. Aedeagus with distinct crest
at base of S-curvation, between lateral lobes (fig. 54).
Aedeagus pore broad and short, oval-shaped (fig. 52).
Measurements: Body length 4: 27.0 mm, 9: 20.9
and 22.4 mm; tegmen length d : 32.0 mm, 9: 31.3 and
31.8 mm; head length d: 2.6 mm, 9: 2.1 and 2.3 mm;
pronotum length d: 2.9 mm, 9: 3.5 and 3.6 mm; me-
sonotum length d: 6.1 mm, 9: 6.1 and 6.6 mm; head
width 3: 5.4 mm, 9: 5.4 and 5.7 mm; width of prono-
tal collar 6: 7.7 mm, ©: 7.8 and 8.4 mm.
Distribution (fig. 2). — The only male known of
this species comes from Sibil, Star Range, in the cen-
tral mountain ranges of New Guinea. Two females,
that probably belong to this species, come from near-
by Feramin.
Etymology. — The species is named in honour of
the geologist C. J. Pigram.
Baeturia daviesi sp. n.
(figs. 2, 60-66)
Type material. — Holotype d : ‘Papua, Mt Lamington, G.
H. Muray, 1924’ (written); ‘W. W. Froggatt Collection’
(print); CANB (print), CSIRO.
Of this species only one male is available. B. davie-
si is very similar to B. pigrami in the shapes of the
male operculum and genitalia, but B. davies: is distin-
guished by a distinctly smaller size, immaculate teg-
mina and the absence of distinct colour markings,
characteristic to B. pigrami and most other species of
1074
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 60-67. — 60-66, Baeturia daviesi n. sp., 60, pygofer in lateral view; 61, pygofer from aslant; 62, operculum; 63, caudo-
dorsal beak in dorsal view; 64, clasper; 65, aedeagus from behind; 66, aedeagus in lateral view. — 67, Baeturia hartonoi sp. n.,
fore femur.
the B. loriae group, though its abdomen is densely
brown speckled, as in B. wegeneri.
Description
Body dull grey-brown with vaguely lighter colou-
red middorsal band, slightly more distinct on prono-
tum. Abdomen 1.4 as long as head and thorax.
Tegmina 1.1 X as long as body length.
Head: Greyish brown, with vaguely darkened spots
between eyes and lateral ocelli. Postclypeus triangu-
larly protruding, 1.5 as wide as long. Postclypeus
angularly swollen ventrally, almost conically protrud-
ing in lateral view (as in fig. 4) Head narrower than
anterior part of pronotum. Vertex very narrow, with
ocelli close together; distance between lateral ocelli
about as wide as frontal ocellus and 0.8 X the distance
between eye and lateral ocellus.
Thorax: Pronotum greyish brown with vaguely
lighter coloured, middorsal band, most conspicuous
over proximal half. Mesonotum brown with a green-
ish tinge, without special colour markings, apart from
two dark spots in front of slightly reddish cruciform
elevation.
Legs: Proximal spine of fore femur rather short,
about as long as distance to middle spine.
18
Tegmina and wings: Hyaline and immaculate,
veins ochraceous. Tegmina with fairly short, square,
8th apical area, and very narrow hyaline border along
hind margins. This border is slightly broader in wing.
Tymbal organ: Tymbal not curved inwards to-
wards distal margin; connecting bar between tymbal
and abdomen almost parallel to body axis. Six trans-
verse sclerotized ridges spanning the tymbal from
dorsal to ventral tymbal margin. Most proximal ridge
narrowing considerably towards the ventral tymbal
margin and only just reaching it. Six short intercalary
ridges seem to form a band across the tymbal. Traces
of bright red between the ridges, especially on dorsal
half of tymbal.
Operculum (fig. 62): Very similar to that of B. pr
grami, but smaller, with dark stains near apex. Basal
part of operculum somewhat quintangular, with its
greatest length at base of lateral margin of distal part,
weakly tapering to distolateral corner and more
strongly tapering to base of meracanthus. Distal part
large, triangle-shaped and slightly domed, curved to
abdomen near its almost pointed apex, and reaching
to about half-lenght abdominal segment 3. Proximal
% of lateral margin slightly convex, distal 4 almost
straight to apex. Distomedial margin concave near
DE Boer: The loriae group of Baeturia
Figs. 68-75. Baeturia hartonoi sp. n. — 68, pygofer in lateral view; 69, caudodorsal beak in dorsal view; 70, clasper; 71, post-
clypeus in lateral view; 72, operculum; 73, aedeagus in lateral view; 74, aedeagus from behind; 75, detail crest between later-
al lobes of aedeagus.
apex and convex to almost rectangular medial corner.
Medial margin short and straight. Opercula close to-
gether medially, separated by narrow and distally lo-
bate 1st sternite. Meracanthus very short, reaching to
about % the operculum length.
Abdomen: Light brown with brown speckling,
darker brown dorsally and lateroventrally, with a
slightly lighter coloured narrow middorsal line,
though without a distinct row of latero-ventral spots.
Segmental hind margins bright red. First tergite only
slightly shorter than 2nd. Anterior margin of 2nd ter-
gite weakly convex middorsally. Ventrolateral part of
2nd tergite, auditory capsules, and anterolateral cor-
ners of 3rd tergite, curved inwards towards ventral
corner of tymbal. Anterolateral part of 2nd tergite ad-
jacent to tymbal.
Genitalia: Pygofer in lateral view (fig. 60) slender.
Dorsal margin straight, continuous with straight,
slender and erect caudodorsal beak. Distal margin of
pygofer straight, angularly bending into margin of
beak. Lateral lobes of pygofer bent inwards, with
small, bluntly rounded protuberances. Caudodorsal
beak very short, pointed at apex (fig. 63). Ventral
margin convex, but weakly concave to base of pygof-
er; ventral margins converging to sharp angle at base
of pygofer opening (fig. 61). Clasper (fig. 64) much
resembling that of B. pigrami, without clasper heel,
but with dorsal margin strongly bent upwards from
clasper base, forming a long finger-shaped, slightly in-
curved protuberance. Dorsal margin strongly bent
outwards distad to protuberance, forming a broad
and distinctly laterally protruding crest. Clasper
slightly bending down to rounded apex. Apical part
of clasper with small clasper hollow. Aedeagus in lat-
eral view (fig. 66) slightly concave along margin of
pore, with narrow and elongate lateral lobes.
Aedeagus with distinct crest at base of S-curvation,
between lateral lobes. Aedeagus pore broad and short,
oval-shaped (fig. 65).
Measurements: Body length: 23.9 mm; tegmen
length: 25.6 mm; head length: 2.0 mm; pronotum
length: 2.8 mm; mesonotum length: 5.1 mm; head
width: 4.8 mm; width of pronotal collar: 6.5 mm.
Distribution (fig. 2). — The only specimen available
comes from Mt. Lamington on the Papuan
Peninsula.
Etymology. — The species is named in honour of
the geologist H.L. Davies.
Baeturia hartonoi sp. n.
(figs. 1, 67-75)
Type material. — Holotype d : ‘PNG: NEW GUINEA: SE: S
Highlands Distr.: Kutubu: Tugiri, 1000 m, 7-9.ii.1978°
(print); ‘J. L. Gressitt Collector BISHOP Museum’ (print),
BPBM.
Of this species only one male specimen is available.
B. hartonoi is a small species, that can be recognized
by an elongate apical part of aedeagus and a lateral
protuberance on the clasper.
19
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Description
Body ochraceous brown, head and thorax slightly
reddish. Abdomen with conspicuous dark dorsal and
midlateral bands, and slightly inflated. Abdomen
1.3X as long as head and thorax. Tegmina 1.3 x as
long as body length.
Head: Brown, slightly darkened around ocelli, but
not as distinctly marked as in some other species of
the B. loriae group. Postclypeus distinctly protruding
beyond vertex lobes, broadly rounded at anterior
margin, and 1.8 X as broad as long. Postclypeus only
very slightly swollen ventrally, anterior margin (later-
al view) weakly convex (fig. 71). Head narrower than
anterior part of pronotum. Distance between lateral
ocelli about 1.5X the width of frontal ocellus and
1.2 X the distance between eye and lateral ocellus.
Thorax: Pronotum, plain brown with no special
markings. Mesonotum almost plain grey-brown, cru-
ciform elevation slightly lighter ochraceous. Two
vaguely darkened spots in front of elevation hardly
visible.
Legs: Proximal spine of fore femur strongly bent,
almost adjacent to femur, and shorter than distance
to middle spine (fig. 67).
Tegmina and wings: Hyaline and immaculate, ve-
nation ochraceous. Tegmen with narrow hyaline bor-
der along hind margin, wing with fairly broad border.
Tymbal organ: Tymbal not curved inwards to-
wards distal margin; connecting bar between tymbal
and abdomen almost parallel to body axis. Six sclero-
tized transverse ridges spanning the tymbal from dor-
sal to ventral tymbal margin. A 7th, most proximal,
ridge almost reaching ventral tymbal margin. Seven
short intercalary ridges seem to form a midlateral
band across tymbal.
Operculum (fig. 72): Short and oblong shaped as
in B. silveri. Basal part of operculum oblong; equally
long at lateral and medial margins. Distal part broad-
er than long, flat against the body. Operculum direct-
ed mesiad, so that connecting bar between tymbal
and abdomen becomes partly visible in ventral view.
Lateral margin very short, bending gradually into
slightly convex distal margin. Distomedial corner an-
gular. Medial margin almost straight. Medial part of
operculum reaching beyond margin of abdominal
segment 2. Opercula widely separated medially, by
broad and rounded Ist sternite. Meracanthus reach-
ing to about 3/4 the operculum length.
Abdomen: Ochraceous, greenish tinged, darkened
in dorsal and lateral bands, though less intensely in
narrow middorsal line. Latero-ventral row of dark
spots most clear on segments 3-5 on right side, and
on segments 3-4 on left side of body, but faded in
successive segments. First tergite very short medially.
Anterior margin of 2nd tergite weakly convex medial-
ly. Lateral parts of 2nd tergite not incurved.
20
Genitalia: Pygofer in lateral view (fig. 68). Dorsal
margin straight and angularly bending into slender
and erect caudodorsal beak. Distal margin broadly
rounded into margin of beak. Lateral lobes of pygofer
strongly bent inwards, with small and bluntly round-
ed protuberances. Ventral margin weakly convex, but
distinctly concave towards base of pygofer. Caudo-
dorsal beak in dorsal view (fig. 69) slender and point-
ed at apex. Clasper very broad, squarely rounded in
lateral view (fig. 70), abruptly broadening dorsally,
distally of clasper base, so that no clasper heel is
formed. Dorsal margin angularly bending down at
about half-lenght, forming a small and slightly in-
curving dorsal protrusion on clasper. This protrusion
seems to function as a support or lock for the aedea-
gus, since it curves inwards, around aedeagus. This in
contrast with the dorsal protrusions on the claspers of
the three foregoing species, which lie proximally of
the aedeagus. Dorsal margin of clasper bending into
almost straight distal margin, at rounded, slightly in-
wards curved, distodorsal corner. Claspers slightly di-
verging towards pointed apices. Apical part of clasper
directed downwards, with narrow clasper hollow.
Lateral side of clasper forming a small and angular, la-
miniform lateral protrusion, unique for this species.
Aedeagus elongate in apical part of its S-curvation
(fig. 71), with narrowly elongate lateral lobes and
truncate apex. Aedeagus with distinct crest between
lateral lobes (fig. 75), crest very low mid-between ba-
sal lobes, and strongly concave (seen from behind
(fig. 74). Aedeagus pore narrow and sharply incised,
truncate at aedeagus apex.
Measurements: Body length: 19.0 mm; tegmen
length: 24.5 mm; head length: 1.8 mm; pronotum
length: 2.5 mm; mesonotum length: 4.3 mm; head
width: 4.3 mm; width of pronotal collar: 5.8 mm.
Distribution (fig. 1). — The only specimen known
comes from Lake Kutubu, in central Papua New
Guinea, just south of the central mountain ranges.
Etymology. — The species is named in honour of
the geologist H.M.S. Hartono.
Baeturia fortuini sp. n.
(figs. 2, 76-87)
Type material. — Holotype d : ‘Coll R.I. Sc. N.B. Papua
New Guinea Morobe pr.: Anguaia, 1800 m, 21.v.1988 (st.
050) J. van Stalle, LG. no: 27363’ (print, orange label),
KBIN. — Paratypes: PAPUA: NEW GUINEA (NE): Bulldog rd.,
60 km S Wau, 2070 m, 22-31.v.1964, J. Sedlacek, 16,
BPBM; Dowalo, W Zenag, 2000 m, 4.ii.1971, J.L. Gressitt,
29, BPBM; Vagau, Herzog Mts., 4000 ft, 4-17.1.1965, M.E.
Bacchus, 1, BMNH; U. Watut SW, 1500 m, 3.v.1968, J.L.
Gressitt, 16, BPBM; same data but 1100-1600 m,
30.iv.1968, 1 4 , BPBM.
Other material. — Mt Missim, 2100 m, 15.11.1968, P.
80
79
DE BOER: The loriae group of Baeturia
83
82
Figs. 76-83. Baeturia fortuini sp. n. — 76, pygofer in lateral view; 77, male operculum; 78, first sternite; 79, clasper, Mt.
Missim; 80, idem, holotype; 81, aedeagus from aslant; 82, aedeagus in lateral view; 83, aedeagus from behind.
Colman, 16, 19, BPBM; Wau, ix,1965, J. Sedlacek, 1°,
BPBM; Wau, Morobe Dist, 1200-1300 m, 14-17.1.1963, J.
Sedlacek, 1 2, BPBM; Wau, Morobe Dist., Mt. Missim, 2000
m, 1.v.1966, O. R. Wilkes, 1d, BPBM.
The most striking character of this species is the
row of bronzed triangular spots along the hind mar-
gins of the tegmina. This characer is also found in B.
loriae, described next. B. fortuini is very similar to B.
loriae, but can be separated from that species by its
larger and triangle-shaped male operculum.
Description
Body of males light brown or greenish tinged, with
dark markings, forming dorsal and lateral bands on
abdomen, and with narrow and light middorsal band
over whole length of body. Females provided with ir-
regular dark patches, and a narrow and light middor-
sal band. Male abdomen 1.4-1.7 as long as head
and thorax, of females 0.9-1.0x. Tegmina of males
1.2-1.4X as long as total body length, of females 1.4-
Nose
Head: Greenish or brown. Vertex and postclypeus
with very long setae, as in B. pigrami. Dark spots on
vertex lobes, between eyes and ocelli, and on lateral
parts of postclypeus. Postclypeus triangularly pro-
truding and 1.4-2.2 x as wide as long. Postclypeus
angularly swollen ventrally (as in fig. 4). Head nar-
rower than anterior part of pronotum. Distance
between lateral ocelli 1.1-1.4X as long as distance
between eye and lateral ocellus.
Thorax: Greenish, with light brown streaks, partly
along oblique fissures, and with two pairs of parame-
dian black spots, at both ends of a broad, greenish
tinged, and immaculate middorsal band. Long setae,
predominantly on these black areas and on pronotal
collar. Colour pattern on mesonotum very variable.
Cruciform elevation and small triangle-shaped area in
front of elevation light brown or bright green, lateral
parts of mesonotum either completely, or only the
distal half, dark brown to castaneous.
Legs: Ochraceous, fore femora with dark brown
streaks. Proximal spine of fore femur shorter than dis-
tance to middle spine.
21
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Fig. 84. Baeturia fortuini sp. n., male tegmen and wing.
Tegmina and wings (fig. 84): Hyaline, though teg-
men with a regular row of seven triangular dark spots,
close to hind margin, with one spot in each apical ar-
ea from apical area 2 to 8. The 8th apical area of teg-
men generally broad and almost square-shaped.
Venation red. Tegmina and wings with fairly broad
hyaline border along hind margins.
Tymbal organ: Tymbal somewhat incurved ven-
trally, connecting bar between abdomen and tymbal
directed inwards. Six transverse sclerotized ridges
spanning the tymbal, from dorsal to ventral tymbal
margin. Most proximal ridge narrowing considerably
towards ventral margin and only just reaching it.
Some specimens with a 7th ridge, hardly separated
from proximal tymbal margin, running from dorsal
margin to about half-width of tymbal. Six short inter-
calary ridges seem to form a midlateral band across
tymbal.
Opercula: Male operculum (fig. 77) ochraceous
with longitudinal dark streaks on distal part. Basal
part of operculum almost oblong, slightly elongate at
half-width and slightly tapering towards medial mar-
gin. Distal part very large, triangular, and slightly
domed, narrowing towards narrowly rounded and al-
most pointed apex, and reaching to about half-length
of abdominal segment 3. Lateral margin of opercu-
lum convex near operculum base, concave at half-
length and convex again near apex. Operculum
strongly curved to body along distomedial margin.
Distomedial margin weakly convex. Medial margin
short and straight. Opercula close together medially,
separated by globularly protruding Ist sternite (fig.
78). Meracanthus very short, reaching to about % the
operculum length. Female operculum (fig. 85) sickle-
22
shaped and erect, with weak crest along its distal mar-
gin. Basal part of female operculum forming a narrow
rim around base of meracanthus.
Abdomen: Male abdomen light brown or greenish.
Dark brown markings forming two dorsal bands, sep-
arated by a narrow and lighter coloured middorsal
line, and lateral bands over abdomen. These bands
are interrupted by lighter, often reddish coloured,
segmental hind margins. Latero-ventral row of dark
spots on segments 3-7 very clear. Ventral side of male
abdomen immaculate, with bright red segmental
hind margins. First tergite quite long and weakly
curved. Anterior margin of 2nd tergite medially
straight. Lateral parts of 2nd tergite hardly inflated,
forming a weak crest along anterolateral margin, au-
ditory capsules somewhat removed from ventral cor-
ner of tymbal. Female abdomen more intensely and
irregularly dark stained. Dark lateral bands less con-
spicuous than in males. Light middorsal line often
very clear, broader than in males. Segmental hind
margins ochraceous or reddened. Ventral side of fe-
male abdomen dark brown stained. Female caudo-
dorsal beak (fig. 86) light ochraceous, long and slen-
der, pointed at apex.
Male genitalia: Pygofer in lateral view (fig. 76)
rather stout, with very long and slender caudodorsal
beak. Dorsal margin concave, gradually bending into
convexly bent beak. Distal margin straight, almost
forming right angle with margin of beak. Lateral
lobes of pygofer strongly curving inwards towards dis-
tal margin, with well-developed, angularly rounded,
protuberances. Ventral margin weakly convex, dis-
tinctly concave near base of pygofer. Caudodorsal
beak in dorsal view (fig. 87) long and slender, point-
ed at apex. Clasper in lateral view (fig. 80) square-
shaped as in B. hartonoi, dorsally abruptly broaden-
ing, distally of clasper base, so that no clasper heel is
formed. Dorsal part of clasper forming a broad and la-
miniform protrusion, broadly rounded at its proximal
corner, and rectangular at its distal corner. Claspers
very narrow and directed downwards towards weakly
diverging apices, and with a narrow and rounded clasp-
er hollow. Both specimens from Mt. Missim with quite
different clasper (fig. 79), more resembling those of B.
silveri and B. loriae, with broad and laminiform, but
less angular, dorsal protrusion and forming a very
small, slightly inwards curved, protrusion at its rectan-
gular distal corner. Aedeagus in lateral view (fig. 82)
very long and slender, with extremely long and slender
lateral lobes, and a very distinct ridge between these
lobes (fig. 81). Aedeagus pore slightly larger than in
foregoing species. Margin of aedeagus pore distinctly
concave at aedeagus apex (fig. 83).
Measurements: Body length d: 22.3-24.5 mm (x
23.4 mm + 0.9), 2: 21.6-22.8 mm (x 22.2 mm # 0.4);
tegmen length 4: 27.0-31.3 mm (x 29.8 mm + 1.5),
2: 31.4-36.6 mm (x 34.6 mm + 1.7); head length 6:
1.7-2.0 mm (x 1.9 mm), 2: 2.1-2.3 mm (x 2.2 mm);
pronotum length d : 2.5-3.0 mm (x 2.8 mm), 9: 3.2-
3.5 mm (x 3.4 mm); mesonotum length d: 4.5-5.4
mm (x 4.9 mm), 9: 5.7-6.4 mm (x 6.1 mm); head
width d: 4.6-5.2 mm (x 5.0 mm), 2: 5.5-5.9 mm (x
5.7 mm); width of pronotal collar d: 6.2-7.4 mm (x
6.8 mm), 2: 7.6-8.2 mm (x 7.9 mm).
Distribution (fig. 2). — B. fortuini seems restricted
to a small area in Papua New Guinea, just south and
west of the Huon Gulf.
Etymology. — The species is named in honour of
the geologist A. R. Fortuin.
Baeturia loriae Distant, 1897 stat. n.
(figs. 2, 88-93)
Baeturia loriae Distant, 1897: 382.
Gymnotympana loriae, Distant 1906: 158; Metcalf 1963:
153.
Identification of types: In his description Distant
(1897) indicates that the type series of this species
contains at least one male and one female, originating
from Moroka and Paumomu river, and collected by
Loria. Two males from Moroka could be traced: one
with a type label was found in the BMNH, the other,
with identification label: ‘Baeturia loriae Dist.’, was
located in the MSNG. Females, or material from
Paumomu river, that could possibly belong to the
type series, were not found. The specimen from the
BMNH is hereby designated lectotype, the other speci-
men belongs to a different species, possibly Baeturia
DE BOER: The loriae group of Baeturia
bicolorata Distant. The lectotype bears the following
labels: ‘Type’ (round label, red margin); ‘loriae Dist
(written); ‘New Guinea SE Moroka, 1300 m, Loria,
vii-x1.93° (print); “Distant coll. 1911. 383’ (print).
Classification. — The reasons for the re-allocation
of this species in the genus Baeturia are given in the
paragraph on phylogeny of the B. loriae group
Only one male of this species is known. B. loriae close-
ly resembles B. fortuint in body markings, and shares the
triangular dark spots in apical areas of tegmina with that
species. B. loriae can be separated from B. fortuini by its
flat and broader, almost rectangular, operculum.
Description
Head and thorax brown, abdomen olive green.
Dark, almost black, markings forming regular dorsal
and lateral bands on abdomen. A narrow and imma-
culate middorsal band over whole length of body.
Abdomen 1.5 X as long as head and thorax. Tegmen
1.2 x as long as total body length.
Head: Grey-brown, with dark brown spots on vertex
lobes and lateral parts of postclypeus. Postclypeus
broadly protruding, 1.8 as wide as long, anterior
margin broadly rounded. Postclypeus distinctly swol-
len and ventrally conically protruding, anterior margin
(lateral view) broadly rounded. Head narrower than
anterior part of pronotum. Distance between lateral
ocelli 1. 4 distance between eye and lateral ocellus.
Thorax: Pronotum ochraceous, with brown streaks
in and along oblique fissures and two pairs of dark
brown spots at both ends of immaculate and lighter
coloured middorsal band. Mesonotum with two large
and triangle-shaped dark spots, in front of cruciform
elevation, and two brown straeks laterad of elevation.
Tegmina and wings: Hyaline, veins ochraceous.
Tegmina, as in B. fortuini. with a regular row of sev-
en triangular dark spots, close to hind margin, one
spot in each apical area from area 2 to 8. The 8th ap-
ical area of tegmen short, almost square-shaped.
Tegmina and wings with a fairly broad hyaline border
along hind margin.
Tymbal organ: Six transverse sclerotized ridges
spanning the tymbal, from dorsal to ventral margin.
Most proximal ridge narrowing considerably towards
ventral tymbal margin and only just reaching it. A 7th
ridge, hardly separated from proximal tymbal margin,
running from dorsal margin to about half width of
tymbal. Six short intercalary ridges seem to form a
midlateral band across tymbal.
Operculum (fig. 91): Ochraceous. Basal part of op-
erculum oblong. Distal part very large and angular,
flat against body. Lateral margin straight, bending
into convex medial margin, at almost rectangular dis-
tal corner. Medial margin bending angularly back to
base of meracanthus, at medial corner. Meracanthus
25
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
93
Figs. 85-93. — 85-87, Baeturia fortuini sp. n., 85, female operculum; 86, female caudodorsal beak in dorsal view; 87, male cau-
dodorsal beak in dorsal view. — 88-93, Baeturia loriae, 88, pygofer in lateral view; 89, clasper; 90, caudodorsal beak in dorsal
view; 91, operculum; 92, aedeagus in lateral view; 93, aedeagus from behind.
very short, reaching to about % the operculum length.
Abdomen: Greenish brown. Dark brown markings
forming two dorsal bands, separated by a lighter nar-
row middorsal line, and lateral bands over abdomen.
These bands are interrupted by lighter, often reddish
coloured, segmental hind margins. Latero-ventral
row of dark spots on segments 3-7 very clear. Ventral
side of male abdomen immaculate, with bright red
segmental hind margins. Anterior margin of 2nd ter-
gite straight medially. Lateral parts of 2nd tergite
hardly inflated, forming a weak crest along anterolat-
eral margin, auditory capsules somewhat removed
from ventral corner of tymbal.
Genitalia: Pygofer in lateral view (fig. 88) slender,
with long and slender, slightly bent, caudodorsal
beak. Dorsal margin of pygofer straight, angularly
bending into beak. Distal margin straight and broad-
ly rounded into margin of beak. Lateral lobes of py-
gofer strongly bent inwards, with bluntly rounded
protuberances. Ventral margin weakly convex, dis-
tinctly concave near base of pygofer. Margins of cau-
dodorsal beak in dorsal view (fig. 90) converging
from base, but parallel in apical part of beak.
Caudodorsal beak rounded at apex. Clasper in lateral
view (fig. 89) angular, square-shaped, as in the Mt.
Missim specimens of B. fortuini. Dorsal margin of
clasper very slightly bending upwards, distally of
24
clasper base, so that no clasper heel is formed.
Straight dorsal margin bending almost rectangularly
into straight distal margin. Distal margin concavely
incurved, just before reaching downwards directed
apical part of clasper. Clasper hollow small and nar-
rowly rounded at clasper apex. Aedeagus in lateral
view (fig. 92) very long and slender, with extremely
long and slender lateral lobes, and a very distinct ridge
between lobes. Aedeagus pore oval (fig. 93).
Measurements: Body length: 24.1 mm; tegmen
length: 30.0 mm; head length: 1.9 mm; pronotum
length: 2.9 mm; mesonotum length: 5.0 mm; head
width: 5.2 mm; width of pronotal collar: 6.9 mm.
Distribution (fig. 2). — The only specimen available
comes from Moroka on the Papuan Peninsula.
Baeturia tenuispina Blöte, 1960
(figs. 1, 94-102)
Baeturia tenuispina Blöte, 1960: 72, figs. 22-23. — Duffels &
Van der Laan 1985: 254.
Material examined. — PAPUA: NEW GUINEA (NE): Gewak,
Salawaket Range, 1530 m, 6.ix.1956, E.J. Ford Jr., 1d, BPBM;
Gurakor, 7.vii.1965, H. Pyka, 18, 72, SMN; Kokoda, 1200
ft, viii.1933, L.E. Cheesman, 16 holotype Baeturia tenuispi-
na Blöte, 36, BMNH; same data, 1d paratype, RMNH; same
98 99
DE Boer: The loriae group of Baeturia
Figs. 94-102. Baeturia tenuispina Blôte. — 94, pygofer in lateral view; 95, pygofer from aslant; 96, male operculum; 97, clasp-
er; 98, male caudodorsal beak in dorsal view; 99, aedeagus from behind; 100, aedeagus in lateral view; 101, female opercu-
lum; 102, female caudodorsal beak in dorsal view.
data but vi-x.1933, 9d, 65 ®, BMNH; Kokoda, xi.1957, 29,
BMNH; Kokoda, 400 m, 15-20.xi.1965, J. and M. Sedlacek,
16, BPBM; Kokoda-Pitoki, 400 m, 25.1ii.1956, J.L. Gressitt,
1d, BPBM; Sangeman Village nr Busu R., NE of Lae, 25 m,
30.viii.1957, D. Elmo Hardy, 34, BPBM; Wau, Morobe
Dist., 1200 m, 29-30.ix.1963, J. Sedlacek, 1d, 12, BPBM; PA-
PUA: NEW GUINEA (SE): Bori nr Sasambota, Popondetta
Subdist., 31.2.1963, D.K. McAlpine, 16, ams; Mt
Lamington Dist, Northern Division, 1925, CT.
McNamara, 19, AMS; same data but vii.1927, 29 ; i-11.1929,
2d, 19, all ams; Popondetta, 25 m, v.1966, Shanahan-
Lippert, 1d, BPBM; same data but vi.1966, 2d, BPBM;
D'ENTRECASTAUX ISLANDS: GOODENOUGH: Goodenough Id.,
x.1943, F/D, C. Ralph, 16, MvM.
Males of B. tenuispina are easily recognized by the
long spine-shaped caudodorsal beak and a small fin-
ger-shaped protrusion on the clasper heel. This pro-
trusion forms the only indication that the species
might be related to the B. loriae group.
Description
Body ochraceous to castaneous brown, covered
with brown speckling. Females on average shorter
than males, but with equally large head and thorax.
Abdomen of males 1.3-1.5 X as long as head and tho-
rax, of females 1.0-1.2 x. Tegmen of males 1.1-1.2 x
as long as body length, of females 1.3-1.4x.
Head: Ochraceous, dark brown or blackish speck-
led. Postclypeus angularly protruding, 1.7-2.3X as
wide as long, anterior margin convex. Postclypeus
slightly swollen ventrally, anterior margin (lateral
view) weakly convex.
Thorax: Pronotum ochraceous brown speckled
medially, between medial pair of oblique fissures.
Mesonotum greyish brown, densely brown speckled
but without any distinct black spots in front of cruci-
form elevation.
Tegmina and wings: Hyaline, venation ochraceous
or reddish, often quite densely set with short setae.
Tegmen with 8 apical areas, the 8th apical area long-
er than in most species of the B. loriae group. No sex-
ual dimorphism in wings. Tegmen with narrow hya-
line border along hind margin, this border distinctly
broader in wing.
Tymbal organs: Tymbal not curved inwards to-
wards distal margin; connecting bar between tymbal
and abdomen almost parallel to body axis. Seven scle-
rotized transverse parallel ridges spanning the tymbal
from dorsal to ventral margin and an 8th, most prox-
imal ridge nearly reaching ventral margin. Seven
short intercalary ridges seem to form a midlateral
band across tymbal. Tymbal reddish coloured at dor-
sal margin and dorsally between ridges.
Opercula: Male operculum (fig. 96) quite large and
almost completely covering tymbal cavity in ventral
view. Distal part of male operculum broad, angularly
rounded, oblong and curved towards body. Lateral
25
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
margin short, convexly bending into long weakly
convex distal margin. Distomedial and medial mar-
gins weakly convex. Medial corners rounded. Meracan-
thus reaching to about 4 operculum length. Female op-
erculum (fig. 101) with very short, sickle-shaped and
erect distal part. Medial margin short and straight
Abdomen: Male abdomen light brown or ochra-
ceous, densely brown speckled dorsally; ventral part,
and sometimes lateral parts of segments 4-5, unspeck-
led. Latero-ventral row of darkened spots generally
very distinct on segments 3-4, but often only vaguely
visible on segments 5-7. Segmental hind margins
bright red. First tergite quite short. Anterior margin
of 2nd tergite medially convex. Lateral parts of 2nd
tergite distinctly swollen adjacent to tymbal. Female
abdomen greyish brown and densely speckled, latero-
ventral row of spots less distinct. Segmental hind
margins often ochraceous. Ovipositor sheaths just
reaching apex of caudodorsal beak. Female caudodor-
sal beak in dorsal view (fig. 102) very long and slen-
der, triangle-shaped and sharply pointed at apex.
Male genitalia: Pygofer (figs. 94-95) very character-
istic, with broadly rounded lateral lobes and long,
slender and erect caudodorsal beak. Dorsal margin
concave to base, but convexly bent into caudodorsal
beak. Distal margin angularly convex, forming a nar-
rowly rounded angle with straight margin of beak.
Ventral margin convex. Lateral lobe of pygofer weak-
ly curving inwards towards end of distal margin,
forming a weakly swollen elongate and bluntly
rounded protuberance. Caudodorsal beak in dorsal
view (fig. 98) extremely long and slender, its parallel
margins converge at about half-length of beak, to-
wards pointed apex. Claspers conspicuous by slender
finger-shaped protuberance on corner of clasper heel
(fig. 97). Dorsal crest very stout and strongly outcurv-
ing, abruptly ending at distolateral corner of clasper.
Apical part of clasper slightly curved down and with
small clasper hollow. Aedeagus (fig. 100) very small
with narrow lateral lobes, in lateral view weakly con-
cave along its pore. Aedeagus pore oval (fig. 99).
Measurements: Body length d: 18.3-23.3 mm (x
20.3 mm + 1.4), 2: 16.7-20.0 mm (x 18.5 mm + 1.2);
tegmen length d: 20.0-24.9 mm (x 22.7 mm + 1.2),
2: 23.6-26.5 mm (x 24.9 mm + 1.2); head length d:
1.6-2.0 mm (x 1.8 mm), 2: 1.6-1.9 mm (x 1.8 mm);
pronotum length d: 2.3-3.2 mm (x 2.7 mm), 9: 2.7-
3.0 mm (x 2.9 mm); mesonotum length d: 4.2-5.4
mm (x 4.6 mm), 2: 4.2-4.9 mm (x 4.6 mm); head
width d: 4.4-5.2 mm (x 4.8 mm), 2: 4.5-5.0 mm (x
4.8 mm); width of pronotal collar d: 5.5-7.1 mm (x
6.3 mm), 2: 6.2-6.9 mm (x 6.6 mm).
Distribution (fig. 1). — B. tenuispina is distributed
in the eastern parts of Papua New Guinea and is also
recorded from Goodenough Island of the D’Entre-
casteaux islands.
26
ACKNOWLEDGEMENTS
For the loan of material I am indebted to: Mr B. J.
Day and Mr M. S. Moulds (Ams); Dr W. J. Knight
and Mr M. D. Webb (BMNH); Mr G. M. Nishida and
Mr K. Arakaki (BPBM); Dr B.P. Moore (CsIRO); Mr J.
van Stalle (KBIN); Dr V. Raineri (MSNG); Dr A.
Neboiss (MVM); Mr J. van Tol (RMNH); and to Mr F.
Heller (SMN).
I would like to thank Mr G. Verlaan for technical
assistance and and Mr D.A. Langerak for the drawing
of figures 8, 9, 59, and 84 and the maps (figs. 1 and
2). I am indebted to Prof. Dr F.R. Schram and Dr
J. P. Duffels (Institute for Systematics and
Population Biology, Amsterdam) for their critical
reading and comments on the manuscript.
REFERENCES
Blôte, H. C., 1960. The genus Baeturia Stâl as represented
in New Guinea (Homoptera, Cicadidae). — Zoologische
Mededelingen 37: 61-80.
Boer, A. J. de, 1982. The taxonomy and biogeography of the
nasuta group of the genus Baeturia Stal, 1866
(Homoptera, Tibicinidae). — Beaufortia 32: 57-78.
Boer, A. J. de, 1986. The taxonomy and biogeography of the
conviva group of the genus Baeturia Stal, 1866
(Homoptera, Tibicinidae). — Beaufortia 36: 167-182.
Boer, A. J. de, 1989. The taxonomy and biogeography of the
bloetei group of the genus Baeturia Stal, 1866
(Homoptera, Tibicinidae). — Beaufortia 39: 1-43.
Boer, A. J. de, 1990. Aedeastria, a new cicada genus from New
Guinea, its phylogeny and biogeography (Homoptera,
Tibicinidae), preceded by a discussion on the taxonomy of
New Guinean Tibicinidae. — Beaufortia 40: 63-72.
Boer, A. J. de, 1991. Scottotympana, a new cicad genus from
New Guinea, with the description of three new species,
their taxonomy and biogeography (Homoptera,
Tibicinidae). — Beaufortia 42: 1-11.
Boer, A. J. de, 1992. The taxonomy and biogeography of the
viridis group of the genus Baeturia Stal, 1866
(Homoptera, Tibicinidae). — Bijdragen tot de Dierkunde
61 (3): 163-183.
Distant, W. L., 1897. Viaggio di Lamberto Loria nella
Papuasia orientale, xvii. Additions to our knowledge of the
Cicadidae of New Guinea. — Annali del Museo civico di
Storia naturale Giacomo Doria Genova (2) 17: 378-383.
Distant, W. L., 1906. A synonymic catalogue of
Homoptera. Part 1. Cicadidae 1906: 1-207 (Trustees
Indian Museum, London).
Duffels, J. P. & P. A. van der Laan, 1985. Catalogue of the
Cicadoidea (Homoptera, Auchenorhyncha) 1956-1980.
— Series Entomologica 34: i-xvi, 1-414.
Metcalf, Z. P., 1963. General catalogue of the Homoptera,
VII. Part 2. Tibicinidae: i-iv, 1-492. North Carolina
State College, Raleigh, N.C.
Receiveid: 2 July 1993
Accepted: 28 July 1993
JAN PATOCKA
Institut fiir Waldökologie der SAW, Zvolen, Slovakia
DIE PUPPEN DER SPANNER MITTELEUROPAS
(LEPIDOPTERA, GEOMETRIDAE): CHARAKTERISTIK
UND BESTIMMUNGSTABELLE DER GATTUNGEN
Patocka, J., 1994. Die Puppen der Spanner Mitteleuropas (Lepidoptera, Geometridae):
Charakteristik und Bestimmungstabelle der Gattungen. — Tijdschrift voor Entomologie, 137:
27-56, figs. 1-227. [ISSN 0040-7496]. Published 15 July 1994.
Characteristics of the pupae of the family Geometridae (Lepidoptera) and an identification key
for its 161 Central European genera are given. The present system of this family is discussed
from the view of pupal characters. The species examined are listed in an appendix.
Institut fiir Waldökologie der SAW, Stúrova 2, 960 53 Zvolen, Slovakia.
Key words. — Geometridae; Pupae; Taxonomy; Key for genera.
Die Familie der Spanner, Geometridae (Lepido-
ptera) ist eine der größten und wichtigsten Schmet-
terlingsfamilien. Viele ihrer Vertreter sind ernste
Schädlinge in Land- und Forstwirtschaft, einige sind
sogar Vorratsschadlinge. Die Mehrzahl der Arten ist
wegen ihrer Gebundenheit an spezifische Standorte
auch vom ökologischen Gesichtspunkt wichtig und
kommt als Indikatoren der bedrohten Umwelt in
Betracht. Die Kenntnis ihrer jiingeren Entwicklugs-
stadien ist fiir viele Fragen unbedingt notwendig; z.
B. für das Studium der Schädlinge, der Prädatoren
und Parasitoide, der Bioindikatoren, Bodenfauna,
verschiedener Synusien usw. Die bisherige Kenntnis
der Puppen der mitteleuropäischen Geometridae ist
liickenhaft. Speziell die Puppen dieser Familie behan-
delt Khotko (1977) und erwähnt 25 Gattungen. In
der vorliegenden Arbeit werden zum Vergleich 161
mitteleuropäische Gattungen, also mehr als 90 % ih-
res Bestandes beriicksichtigt. An Khotko’s Arbeit
knüpfen die Verôffentlichungen des Verfassers an,
die zahlreiche weitere Arten der Unterfamilie Laren-
tiinae (Patocka 1980-1994) und mehrere Gruppen
der Ennominae (Patoëka 1978, 1985, 1986 a, b,
1992, 1993) behandeln. Auf diese Weise ist zwar die
Mehrzahl der Arten dieser beiden Unterfamilien ein-
gehend bearbeitet, es fehlt jedoch eine zusammenfas-
sende Bestimmungstabelle der Gattungen. Diese
Liicke soll die vorliegende Arbeit ausfiillen. Einge-
hendere Angaben iiber viele Geometridenpuppen fin-
den sich bei Ljungdahl (1919) und Nordström et al.
(1941), über die Arten an Obstbäumen Speyer (1958),
an Tannen Patoëka et al. (1960), an Eichen Patocka
(1980) und iiber die Puppen in der Bodenstreu in
Kiefernwaldern Koehler (1937). Einzelbeschreibun-
gen von Geometridenpuppen sind in weiteren Litera-
turquellen verstreut. Die Handbücher über die
Schmetterlinge Mitteleuropas, wie Forster & Wohl-
fahrt (1981) geben nur kurze und für eine Bestim-
mung nicht ausreichende Puppenbeschreibungen.
Die Puppen der kanadischen Geometridae wurden
im Rahmen einer Monographie dieser Familie von
McGuffin (1967-1981) bearbeitet.
Das hier beniitzte System und die Nomenklatur
folgen grundsatzlich Leraut (1980) und Gustafsson
(1987), die morphologische Terminologie richtet
sich nach Mosher (1916) und McGuffin (1967-
1981). Das untersuchte Material (die hier beriick-
sichtigten Gattungen und Arten sind im Anhang auf-
gelistet) stammt grofsenteils aus der Sammlung des
Verfassers, zu welcher viele Fachkollegen durch
Uberlassung von Puppen und Puppenexuvien beige-
tragen haben, wofür ihnen auch an dieser Stelle herz-
lich gedankt sei. Viele Arten konnte sich der Verfasser
von Museen ausleihen, insbesondere von dem
Museum für Naturkunde der Humboldt-Universität
in Berlin (Dr. W. Mey), von dem Naturhistorischen
Museum Wien (Dr. M. Lédl), von dem Zoologi-
schen Museum Kopenhagen (O. Karsholt), von der
Zoologischen Staatssammlung Miinchen (Dr. W.
Dierl) und aus der Entomologischen Abteilung des
Nationalmuseums in Prag (Dr. F. Krampl). Den ge-
nannten Institutionen und Herren dankt der Ver-
fasser verbindlichst, ebenso wie auch der Alexander
von Humboldt-Stiftung Bonn fiir die Erteilung eines
Stipendiums, welches ihm das Studium des Materials
in München, sowie der Hanskarl-Göttling-Stiftung
der Bayerischen forstlichen Versuchs- und For-
schungsanstalt, welche einen Aufenthalt in Berlin er-
möglichten.
27
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Die Puppen und ihre Exuvien wurden mit Hilfe ei-
nes Stereomikroskopes und z. T. eines Abbeschen
Zeichenapparates untersucht und gezeichnet.
CHARAKTERISTIK DER FAMILIE GEOMETRIDAE
Eine Ubersicht der Puppenmorphologie dieser
Familie geben die Abb. 1-6. Die Puppen der mittel-
europäischen Geometridae sind kleine bis mittel-
große (etwa 5-25 mm lang) Pupae obtectae. Sie sind
schlank (Abb. 86) bis gedrungen (Abb. 94), meistens
in der Mitte am breitesten, vorne abgerundet und
hinten mehr zugespitzt (Abb. 1, 2, 88, 94). Manche
Formen (z.B. Cyclophora, Abb. 130) sind vorne durch
einen Höcker an der Basis der Vorderflügel breiter
und sehen in Ventralansicht eher keilartig aus. Die
Mehrzahl der Puppen ist einfarbig gelb-, rot-, oder
schwarzbraun bis fast schwarz gefärbt und + glän-
zend. Es handelt sich hierbei um Arten die sich im
Boden oder in der Bodenstreu, in einem Erdkokon
bzw. einer Erdhöhle verpuppen. Die übrigen Puppen
sind griin, gelblich oder sandfarben, + glanzlos und
oft mit dunklen Zeichnungen versehen. Das sind
Arten, die sich oberirdisch bzw. am Boden in einem
leichten Gespinst, oder nur mit Einzelfaden befestigt
verpuppen. Eine Gattung (Abraxas) ist glänzend
schwarz mit satt gelben Querstreifen am Abdomen,
die sich ebenfalls oberirdisch verpuppt.
Die Oberflache der Puppen ist fein bis grob skulp-
turiert. Meist ist der Kopf und Thorax + gerunzelt
oder gefurcht und das Abdomen (manchmal auch der
Thorax, insbesondere das Metanotum) mit kleinen
punktförmigen Griibchen, den sog. Punktgriibchen
(Abb. 210) besetzt. Selten entspringen von diesen
Microbörstchen (Apeira, Abb. 74). Bei wenigen
Puppen (z. B. bei der Gattung Geometra, Abb. 78)
kommt eine Stachelskulptur vor. Die Körperborsten
sind kaum sichtbar bis relativ groß und stark (Abb.
31). Der Vertex wächst entweder mit der Frons ohne
eine Epicranialnaht zusammen (Vertreter der
Unterfamilie Ennominae und einige andere, Abb. 4),
oder er ist durch eine deutliche Epicranialnaht abge-
trennt (Mehrzahl der übrigen Unterfamilien, Abb. 3).
Beim Schlüpfen löst sich der Vertex bei dieser
Gruppe von der Frons ab. Die Frons ist in wenigen
Fällen mit einem Paar Höcker (Abb. 8, 9, 86, 125)
versehen, oder kegelförmig vorgezogen (Abb. 43), an-
sonst meist abgerundet. Zwei Paare von Frontal-
(Abb. 4) und ein Paar von Clypealborsten (Abb. 116)
sind meist deutlich und groß ausgebildet. Von der
Frontoclypealnaht sind nur Teile nahe der Antennen
deutlich erkennbar. Die Komplexaugen sind groß
und bestehen aus einem + glatten, ventralen Streifen
und einem oft mehr rauhen Dorsalteil (Abb. 95, 96).
Die Mandibeln sind meist kaum differenziert (Abb.
112, 116), bei Theria jedoch stark hervortretend
28
(Abb. 22). Das Labrum erscheint meist halbkreis-
(Abb. 46), schaufel- (Abb. 117) oder trapezförmig
(Abb. 47), gemeinsam mit dem schwach abgegrenz-
ten Postclypeus oft fünfeckig (Abb. 116). Das
Labium ist entweder ganz verdeckt (Abb. 87) oder
klein (Abb. 209) bis groß (Abb. 101), als eine drei-
oder fünfeckige Platte sichtbar. Die Maxillar- und
Labialpalpen sind stets verdeckt. Die Genae grenzen
an die Vorderbeine; diese Grenze immer kürzer als
die Grenze zwischen Auge und Vorderbein. Die
Grenzen zwischen Vorderbeine und Antennen kurz
oder lang (Abb. 19, 21). Bei den sonst ähnlichen
Puppen der Herminiinae, Noctuidae, grenzen die
Vorderbeine nicht an die Antennen, weil die
Mittelbeine die Augen berühren. Die Vorder- und
Mittelbeine sind meist relativ lang, die Vorderbeine
zuweilen (Abb. 16), die Mittelbeine meistens unge-
fähr so lang wie die Antennen und Vorderflügel. Die
Enden der Hinterbeine sind oft sichtbar. Die
Vorderschenkel sind sichtbar, groß (Abb. 1, 125)
oder klein, schmal (Abb. 94, 77, 165), oft aber ganz
verdeckt (Abb. 16, 94a). Die Proboscis ist meist so
lang oder wenig kürzer als die Vorderflügel, selten
stark verkürzt und deutlich kürzer als die Mittelbeine,
bzw. auch als die Vorderbeine, (Abb. 16, 98, 99). In
wenigen Fällen überragt die Proboscis (Abb. 9, 12,
94a, 123) und noch seltener überragen auch die
Mittelbeine und Antennen (Abb. 13) die Spitzen der
Vorderflügel und den Kaudalrand des 4. Abdominal-
segmentes. Bei Rhodometra (Abb. 123) verschiebt die
vergrößerte Proboscis das Labrum bis nahe am
Frontalende des Kopfes, was oft auch bei Sphingidae
mit langer Proboscis vorkommt. Die Antennen sind
meist ungefähr so lang wie die Vorderflügel (Abb. 1).
Die Vorderflügel überragen den Kaudalrand des 4.
Abdominalsegmentes nicht, sind voneinander ge-
trennt (Ausnahme: Apeira, Abb. 76) und zeichnen
sich zuweilen durch hervorstehende (Abb. 94) oder
dunkel gestreifte bzw. helle Adern aus. Selten liegt an
ihren Wurzeln je ein Höcker (Abb. 123, 125, 130).
Die Hinterflügel säumen die Vorderflügel meist zu 4
bis 4 des 4. Abdominalsegmentes, nicht aber bis zu
deren Spitzen, wie es bei der Familie Drepanidae, die
den Geometridae sonst ähnlich ist, vorkommt. Eine
Ausnahme ist die Gattung Archiearis.
Die thorakalen Spiracula sind klein und spaltför-
mig. Oft bildet sich kaudal von ihnen am Mesono-
tum, ein auffälliger, tomentöser (Abb. 31, 91), rauh
skulpturierter (Abb. 54) oder ‘ohrartig hervortreten-
der, kantiger Höcker (Abb. 33, 35). Das Metanotum
besitzt frontal in der Mitte meist einen mitteltiefen
und abgerundeten Ausschnitt. Die Frontolaterale
Lappen des Metanotums sind meist abgerundet
(Abb. 102, 103), zuweilen spitz oder winkelig (Abb.
153), insbesondere bei den Vertretern der Unterfami-
lie Sterrhinae. Die Vorderecken des 1. Abdominalseg-
mentes sind meist abgerundet, Ihre Seiten + parallel.
Die Spiracula sind am 2.-3. Abdominalsegment sicht-
bar oder + verdeckt. Die abdominalen Spiracula sind
klein bis mittelgroR, meist schmal elliptisch, oft mit
breiten elliptischen Höfen und nicht selten + erha-
ben. Die 5.-7. Abdominalsegmente sind beweglich.
An der Basis des 5. Segmentes frontal des Spiracu-
lums finden sich oft Leisten, Gruben und verdichtete
Punktgriibchen (Abb. 34, 38, 40, 45, 49-55, 138,
140-142), die gute Unterscheidungsmerkmale bie-
ten. An der intakten Puppe sind sie jedoch durch den
Kaudalrand des 4. Abdominalsegmentes oft verdeckt;
dieser sollte deshalb, um eine Untersuchung zu er-
möglichen, mit einer Nadel beseitigt werden. An der
Puppenexuvie sind diese Gebilde auch ohne diesen
Eingriff meist gut sichtbar. Bei der Gattung Rheuma-
ptera gibt es Querleisten und Rinnen an der Basis der
Dorsalseite des 5. Abdominalsegmentes (Abb. 139).
Zwischen dem 9. und 10. Segment besitzt die Mehr-
zahl der Spannerpuppen einen + tiefen, rinnenartigen
Quereinschnitt, die sog. Dorsalrinne, welche an ihrer
Kaudalseite meist einen oder mehrere Ausläufer auf-
weist, deren Ränder oft verdunkelt und + tomentös
sind. Seltener bildet die Dorsalrinne auch einen oder
einige Ausläufer frontalwärts (Frontalausläufer, Abb.
6). An den Seiten entspringt von dieser Rinne oft
noch ein dorso-kaudalwärts gerichteter Lateralein-
schnitt (Abb. 5).
Dorsalseite des 10. Abdominalsegmentes ist ver-
schieden lang (Abb. 70, 71), seine Ventralseite oft +
breit mit gewölbten Seiten, abgeflacht, und der
manchmal erhabene frontolaterale Teil durch flache
Furchen abgegrenzt. Die Analnaht ist meist lang, ihre
oft erhabenen Seiten wallartig, manchmal auch mit
Längs-, seltener Querrunzelung. Das sog. Analfeld
grof und auffällig. Die Genitalnaht und ihr Genital-
feld, besonders bei den Männchen sind oft groß. Der
Absturz zur Basis des Kremasters verläuft steil bis
schräg (Abb. 137, 182). Der Kaudalausläufer des 10.
Segmentes, der Kremaster, welcher bei der Puppe ei-
ne Haft- und Stützfunktion besitzt (Patocka 1978),
ist bei den Puppen der Geometridae immer vorhan-
den und + groß, seltener klein (Abb. 17) und mehr
oder weniger scharf von dem 10. Abdominalsegment
abgegrenzt bzw. abgesetzt. In der Dorsalsicht ist er oft
haufen- (Abb. 119) oder zungenförmig (Abb. 80),
dreieckig (Abb. 178), seltener trapez- (Abb. 18, 164)
oder ankerförmig (Abb. 7). Ventral an seiner Basis
gibt es oft ein Paar Basalhöcker, dazwischen eine
Vertiefung bzw. Mulde. Bei der Unterfamilie Geo-
metrinae ist der Kremaster ventral charakteristisch ra-
dial gefurcht (Abb. 75, 79), ansonst ist er + glatt oder
verschiedenartig skulpturiert (gerunzelt, gefurcht, ge-
rippt; Abb. 120, 122). Der Kremaster tragt meist 4,
seltener 1-3 oder 5-7 (bei Apeira eine Anzahl) Paare
von Borsten, die oft am Ende hakenartig gekrümmt
Patoëka: Puppen der Spanner Mitteleuropas
sind. Die Bezeichnung von 4 Paaren von Primärbor-
sten am Kremaster (nach McGuffin 1967-1981) zei-
gen die Abb. 5-6. Bei vielen Geometriden-Puppen
wachsen die starken Borsten D, zu einem Stiel zusam-
men und nur ihre Spitzen bleiben als eine Gabelspitze
frei (Abb. 32), die manchmal auch vermehrt sind
(Abb. 28). Während bei vielen Vertretern der Unter-
familie Larentiinae die übrigen Borsten noch erhalten
bleiben (Abb. 177, 178) verschwinden sie bei zahlrei-
chen Ennominae vollständig (Abb. 71). Bei Bupalus
bleibt nur eine einfache, stumpfe Spitze übrig (Abb.
42).
Die Puppen der Spanner findet man z. T. an der
Nahrungspflanze, durch die Häkchen am Kremaster
und einen Gürtelfaden befestigt (z.B. bei Cyclophora)
oder in einem lockeren Gespinst (Abraxas, Ennomos,
bei den Comibaenini in dem Sack der Raupe) direkt
an der Nahrungspflanze oder auch am Boden. Die
Mehrzahl der Puppen ruht jedoch in einem + festen
Erdkokon oder einer Erdhöhle in der Bodenstreu
oder im Boden selbst. Die Puppen variieren in der
Größe insbesondere nach Geschlecht, Generation
oder relativer Abundanz. Die Raupen leben an ver-
schiedenen Gruppen von Pflanzen, sogar an Sporen-
pflanzen bzw. an Detritus. Es gibt ebenso polyphage,
wie relativ streng oligophage, an spezielle Pflanzen-
gattungen gebundene Arten (z. B. Clematis, Actaea,
Berberis, Rhamnus, Hypericum, Veratrum). Manche
Gruppen (Eupitheciini, Perizomini) bevorzugen
Blüten und Früchte. Viele Arten leben an Laubhöl-
zern, an denen sie oft auch schädlich werden, aber
auch an Nadelhölzern, und eine zahlreiche Gruppe
von Larentiinae lebt an Galium.
Die Spanner bewohnen verschiedenartige Biotope,
sehr viele Wälder und steppenartige, mehrere auch
ruderale Standorte, Siimpfe bzw. Torfmoore, Felsen
und Alpenwiesen im Hochgebirge. Viele Arten sind
fiir bestimmte Biotope recht charakteristisch.
BESTIMMUNGSTABELLE DER GATTUNGEN
Einige Gattungen, wie Orthostixis Hiibner, 1822,
Costaconvexa Agenjo, 1949, Coenocalpe Hiibner, 1825,
Bichroma Gumppenberg, 1887, Eilicrinia Hübner, 1825,
Mannia Prout, 1915, konnten infolge Materialmangels
nicht beriicksichtigt werden.
1 Kaudalhälfte des Kremasters in Dorsalsicht ein-
fach zugespitzt (selten mit zwei Börstchen am
Ende) (Abb A) eee ene Bupalus Leach
— Kaudalhälfte des Kremasters in Dorsalsicht mit
2-4 Dornen oder Zähnen, jedoch ohne
Börstchen oder Häkchen (Abb. 7, 14, 20). Die
Dornen oft gestielt (Abb. 32, 52), zuweilen auch
verzweigt Abb NS) RE Eeen ee À
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
30
= Kaudalhälfte des Kremasters mit 2-8 (seltener
mehr) Häkchen oder Borsten (Abb. 6, 72,
80, 89, 136). Die Endborsten (D,) zuweilen
+ dornartig umgebildet (Abb. 89, 143, 218).
SRO) e len enn EEEN 3
- Frons ohne ein Höckerpaar, selten kegelför-
mig vorgezogen (Abb. 43), sonst abgerundet
(AS, U) eee ER 4
Proboscis überragt deutlich den Kaudalrand
des 4. Abdominalsegmentes (Abb. 9) … … ….
Er Siona Duponchel
= Proboscis überragt den Kaudalrand des 4.
Abdominalsegmentes nicht (vgl. Abb. 1) ......
ee LINO i Aethalura McDunnough
Einzelne Abdominalsegmente dorsal mit gro-
ßen, elliptischen, dunkel umrandeten Gru-
DEN ADD) a else 5
= Abdominalsegmente ohne solche Gruben
(ll Abbi RPM re ru 6
Am 5.-7. Abdominalsegment dorsal fehlen
die Gruben... Lithostege Hübner
= Auch am 5.-7. Abdominalsegment gibt es
dunkel umrandete Gruben (Abb. 11) ...........
Betori ee Chesias Treitschke
Proboscis überragt den Kaudalrand des 4.
Abdominalsegmentes (Abb. 12, 13)........... 7
= Proboscis überragt nicht den Kaudalrand des
4. Abdominalsegmentes (Abb. 1)................ 9
Sowohl die Antennen wie auch Mittelbeine
iiberragen den Kaudalrand des 4. Abdomi-
Mal SCP IMCMTES ne eens tees 8
= Nur die Antennen, nicht jedoch die Mittel-
beine tiberragen den Kaudalrand des 4. Ab-
dominalsegmentes............. Odezia Boisduval
— Weder Antennen noch Mittelbeine überra-
gen den Kaudalrand des 4. Abdominalseg-
Imentesi (Abb MO) Schistostege Hiibner
Kremaster länger als basal breit (Abb. 15).....
RIA Pan ane nre Carsia Hübner
= Kremaster kürzer als basal breit (Abb. 13, 14)
ET Aplocera Stephens
Patoka: Puppen der Spanner Mitteleuropas
9 (6) Kremaster ankerartig, stumpf abgerundet, an
seinen Seiten je ein größerer und oft auch
kleinerer Zahn (Abb. 7). Puppe schlank........
RE ER Ooh Archiearis Hübner
= Kremaster am Ende mit gabelartigen, schrag
kaudalwärts gerichteten Dornen (Abb. 25,
32, 52), diese zuweilen weiter verzweigt
(Abb. 28). Wenn die Dornen + gegeneinan-
der stehen (Abb. 17, 20), nur in einem Paar
und die Puppe ist gedrungen … … 10
Weder Labium, noch Vorderschenkel sicht-
barg(AbbICRI Pen 11
10 (9)
— Labium, Vorderschenkel, oder beides sicht-
bare MAD EEE En 23
11 (10) 8. Abdominalsegment dorsal mit einem Paar
spitzerblocker nr Nychiodes Lederer
= 8. Abdominalsegment dorsal ohne Höcker …
ee KOE ENE SENDEN AR TER 12
12 (11) Kremaster mit zwei Paaren schlanker, wenig
divergierender Dornen, außerdem in seinem
Basalteil noch ein Paar Höcker (Abb. 23) .....
teo: kat EREN PRES Artiora Meyrick
= Kremaster mit 1 Paar Dornen, die zuweilen
noch verzweigt sein können, außerdem höch-
stens mit kurzen Zähnen an den Seiten (Abb.
1952022 HP DR EEEN 13
13 (12) Kremaster klein, kurz, seine Dornen stark di-
vergierend (Abb) 14
= Kremaster groß, am Ende mit langen Dornen
oder mit einer gestielten Gabel (Abb. 25, 27,
14 (13) Proboscis kiirzer als die Vorderbeine (Abb.
lea Operophtera Hübner
— Proboscis länger als die Vorderbeine (Abb. 1)
EE Bento ae 15
15 (14) Grenzen zwischen Vorderbeine und Anten-
nen etwa 2X so lang wie die Grenzen zwi-
schen Vorderbeine und Augen (Abb. 21).
Mandibeln erhaben .............. Theria Hübner
= Grenzen zwischen Vorderbeine und Anten-
nen höchstens wenig langer (oft kürzer) als
die Grenze mit den Augen (Abb. 19). Mandi-
beln nicht erhaben............ Alsophila Hübner
Abb. 1-21. — 1,2,3,5,6, Horisme vitalbata; 4, Synopsia sociaria, 7, Archiearis notha; 8, Aethalura punctulata; 9, 10, Siona line-
ata; 11, Chesias legatella; 12, Schistostege treitschkei, 13, 14, Aplocera praeformata; 15, Carsia sororiata, 16, 17, Operophthera
brumata; 18, 19, Alsophila quadripunctaria; 20, 21, Theria rupicapraria.
1, Habitusbild der Puppe, Ventral; 2, Dorsalsicht; 3, 4, Frons und Umgebung; 5, Abdominalende in Lateralsicht; 6, idem, in
Dorsalsicht; 7, Kremaster in Dorsalsicht; 8, Kopf in Ventralsicht; 9, Frontalhalfte der Puppe in Ventralsicht; 10, 14, 15, 17,
18, 20, Abdominalende in Dorsalsicht; 11, Abdominalhalfte der Puppe in Dorsalsicht; 12, 13, idem in Ventralsicht; 16, Kopf
und Beine in Ventralsicht; 19, 21, Augen und Umgebung.
A,-A,, = 1-10. Abdominalsegment, Af = Analfeld, An = Analnaht, Ant = Antennen, As = abdominale Spiracula, C1 =
Clypeus, D,, D, = Dorsalborsten 1,2, Dr = Dorsalrinne, Es = Epicranialnaht, Fa = Frontalausläufer der Dorsalrinne, Fr =
Frons, G = Genae, Gn = Genitalnaht, Hb = Hinterbeine, Hfl = Hinterfliigel, Ka = Kaudalausläufer der Dorsalrinne, Kr =
Kremaster, L, = Lateralborste 1, Lb = Labium, Lbr = Labrum, Le = Lateraleinschnitt, Mb = Mittelbeine, Msn = Mesonotum,
Mtn = Metanotum, O = Auge, Pcl = Postclypeus, Pn = Pronotum, Pr = Proboscis, Sd, = Subdorsalborste 1, Ts = thorakales
Spiraculum, V = Vertex, Vb = Vorderbeine, Vfl = Vorderfliigel, Vs = Vorderschenkel
31
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
32
16 (13) 9. Abdominalsegment mit einer Querreihe
von Dornen (Abb. 24).. Eurrhanthis Hiibner
= 9. Abdominalsegment ohne Dornen (Abb.
SO) ene IERIE SANT. one ai 17
17 (16) 10. Abdominalsegment mit je einem spitzen
lateralen Fortsatz (manchmal ein weiterer an
der Mitte des Kremasters, Abb. 25) ..............
AI AR bedene Lycia Hübner
- 10. Abdominalsegment ohne, Kremaster oft
mit einem Paar von laterale Fortsätze (Abb.
18 (17) Dorsalrinne vorhanden, Kaudalausläufer +
tomentôs, Lateraleinschnitt deutlich (Abb.
Dr Ds DAS) EEE ARRETE cento negli Paoli 19
= Dorsalrinne und Lateraleinschnitt nicht vor-
ranclens (ADD 67) EEE 21
19 (18) Erhebung kaudal des thorakalen Spiraculums
flach, länglich elliptisch, stark tomentös
(Abb. 31). Puppe 12-15 mm lang..................
ARTE IA Erannis Hübner
= Erhebung kaudal des thorakalen Spiraculums
‘ohrartig’, kantig (Abb. 33, 35). Puppe über
N ee ee 20
20 (19) Leiste frontal des Spiraculums am 5. Abdo-
minalsegment doppelt (Abb. 34). Erhebung
kaudal des thorakalen Spiraculums steiler
frontal- als kaudalwärts (Abb. 33). Puppe 17
MMA. enter. ee. Apocheima Hübner
— Leiste frontal des Spiraculums am 5. Abdo-
minalsegment einfach (Abb. 36). Erhebung
kaudal des thorakalen Spiraculums weniger
steil frontal- als kaudalwärts. (Abb. 35) Pup-
po2 0-2 Ormimilang en: Biston Leach
21 (18) Puppe 17-21 mm lang.. Phigalia Duponchel
— Buppaal0-1Simmllans.e. wen. ee 22
22 (21) Kremaster mit deutlichen lateralen Fort-
sätzen (Abb. 41). Vertiefung frontal des
Spiraculums am 5. Abdominalsegment mit
zahlreichen Punktgrübchen frontal der
Querleiste (Abb. 40)....... Selidosema Hübner
= Kremaster ohne, bzw. nur mit angedeuteten
lateralen Fortsätzen (Abb. 32). Vertiefung
am 5. Abdominalsegment frontal des Spira-
culums mit Querleisten, ohne Punktgriib-
chen (Abb. 37, 38) ............ Agriopis Hübner
23 (10) Dorsalrinne mit 1-2 großen Kaudalausläu-
Patotka: Puppen der Spanner Mitteleuropas
ferni(Abb926 27029) Mine Ren 24
= Dorsalrinne mit mehreren Kaudalauslaufern,
oder nicht vorhanden (Abb. 56, 62, 70) .. 28
24 (23) Vorderschenkel nicht sichtbar (vgl. Abb. 82)
= Vorderschenkel sichtbar (vgl. Abb. 1) … 26
25 (24) Dorsalrinne mit nur einem großen Kaudal-
ausläufer. Kremaster ohne laterale Fortsätze
(AD DR eee Chondrosoma Anker
— Dorsalrinne mit zwei großen Kaudalausläu-
tern (ADD 0277) SA Lignyoptera Lederer
26 (24) Frons breit kegelförmig vorgezogen (Abb.
43). Dorsalrinne auch an der Frontalseite
ausgebuchtet (Abb. 44) … … Pelurga Hübner
= Frons nicht vorgezogen. Dorsalrinne nur an
der Kaudalseite mit einem großen, spitzen
Vorsprunp{(Abb29) Fee 27
27 (26) Puppe mehr gedrungen, schwärzlich, Exuvie
schwarzbraun. Puppenlänge 6-8 mm ............
EREN ee ly Phibalapteryx Stephens
= Puppe schlanker, bräunlich, Exuvie gelb-
braun. Puppenlänge 9-10 mm ......................
BR: ER RN Cataclysme Hübner
28 (23) Dornen am Kremaster voneinander entfernt
entspringendulAbb 30) m een 29
— Dornen am Kremaster gabelartig, gestielt
(ADD PE NC 0252.06) eens eee 30
29 (28) Kremaster an der Dorsalseite fein skulptu-
Hert. huppe nur: iMm lange eters enne
DCO, i ae ferry eh Mee toe Pygmaena Boisduval
— Kremaster an der Dorsalseite grob skulptu-
riertsbuppelüberg Ofman Re
À Die CB EI der Beau Ei Gnophos Treitschke s. lat.
30 (28) Proboscis wesentlich kürzer als die Mittel-
beine. Labium sehr klein. Puppe 7-9 mm
Rings use Tephronia Hübner
= Proboscis nicht oder wenig kiirzer als die
Mittelbeine (im Zweifelsfall Labium Mittel-
ROS) ie MEL PRE a EME AR PE 31
31 (30) Dornen am Kremaster + länger als ihr Stiel
(Abb. 102, 206). Puppen nur 6-8 lang... 152
= Dornen am Kremaster + kiirzer als ihr Stiel
(Abb. 52, 62, 67). Meist größere Puppen .....
Se és A SSS REES a aeons 33
Abb. 22-57. — 22, 39, Theria rupicapraria, 23, Artiora evonymaria, 24, Eurranthis plumistaria, 25, Lycia zonaria, 26,
Chondrosoma fiduciaria, 27, Lignyoptera thaumastaria, 28, 31, Erannis defoliara, 29, Phibalapteryx virgata, 30, Gnophos ob-
fuscatus, 32, 38, Agriopis aurantiaria, 33, 34, Apocheima hispidaria, 35, 36, Biston strataria, 37, Agriopis bajaria, 40, 41,
Selidosema plumaria; 42, Bupalus piniarius, 43, 44, Pelurga comitata; 45, 46, Ectropis crepuscularia, 47-49, Calospilos sylvata;
50, Cleora cinctaria; 51, Fagivorina arenaria, 52-54, Serraca punctinalis, 55, Ematurga atomaria, 56, 57, Chiasmia clathrata.
22, Labrum, Mandibulae; 23-30, 32, 41, 42, 44, 52, 56, Abdominalende in Dorsalsicht; 31, 33, 35, Erhebung kaudal des
thorakalen Spiraculums; 34, 36, 38, Basis des 5. Abdominalsegmentes, Lateralsicht; 29, Kopf und Beine in Ventralsicht; 40,
45, 49-51, 53, 57, Basis des 5. Abdominalsegmentes, Lateralsicht; 43, Kopf in Ventralsicht; 46, 47, Labrum, Labium; 48,
Abdominalende in Lateralsicht; 54, Erhebung kaudal des thorakalen Spiraculums.
55)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
34
= Dorsalrinne nicht entwickelt (Abb. 70, 71)
Pee Ne hie nti ters. A Dl aves 47
33 (32) Kaudal des thorakalen Spiraculums je eine re-
lativ große, breite, sehr rauhe, erhöhte Fläche
(Abb. 54). An der Basis des 5. Abdominalseg-
mentes lateral zwischen zwei dunklen Quer-
leisten wenigstens 12 Zellen in einer Reihe
(AIDES) SRE CR Re PRE 34
— Kaudal des thorakalen Spiraculums eine klei-
ne, längliche oder eine unauffällige Erhebung
(vgl. Abb. 106, 111). Wenn diese jedoch grö-
fer und rauher ist (Abb. 227) und auch
sonst, gibt es an der Basis des 5. Abdominal-
segmentes lateral anders aussehende Bildun-
gen (Abb. 51, 57, 65, 68, 69) oder nur etwa
SACIIEnCAbD 50) PEER LAN re 36
34 (33) Mesonotum ohne Punktgrübchen … … … …
So Base eenn Tes Hypomecis Hiibner
Mesonotum mit Punktgrübchen.............. 35
35 (34) Spiraculum am 5. Abdominalsegment liegt
in der Nähe der 10.-15. Zelle (von der Ven-
tralseite gezählt, Abb. 53). Puppe meist über
14 mm lang, eher glänzend ... Serraca Moore
= Spiraculum am 5. Abdominalsegment liegt
in der Nahe der 2.-6. Zelle (Abb. 55). Puppe
imeistmattery unter U3) laine). ito
per chat dt nel Rut arn ged Ematurga Lederer
36 (33) Abdomen ohne Punktgriibchen (vgl. Abb.
DKM er Lomaspilis Hübner
= Abdomen mit Punktgrübchen (vgl. Abb. 51)
IT o ai gee M ea a 37
37 (36) Am 5. Abdominalsegment frontal des Spira-
culum eine langlich ovale, von einer dunklen
Leiste umgebene Fläche (Abb. 45) … … … 38
= Am 5. Abdominalsegment vor dem Spiracu-
lum andere Bildungen: Zellen (Abb. 50),
Leisten (Abb. 65, 68, 69) oder nur Punkt-
grübchen (Abb. 57) bzw. eine glatte Fläche
CAS re nn 39
38 (37) Labium ganz klein (Abb. 46) bzw. fehlend.
Vorderschenkel gut sichtbar (vgl. Abb. 1).
Lateraleinschnitt undeutlich .........................
RS e ent lee rs Ectropis Hübner
= Labium (Abb. 47) und Lateraleinschnitt
(Abb. 48) deutlich. Vorderschenkel verdeckt
(ele Ab bi39) Eer Calospilos Hübner
39 (37) Frontal des Spiraculums am 5. Abdominal-
Patotka: Puppen der Spanner Mitteleuropas
segment nur eine glatte Fläche, weder mit
Leisten, noch Punktgrübchen (Abb. 51) … …
SEARS A NER Fagivorina Wehrli
— Frontal des Spiraculums am 5. Abdominal-
segment eine dunkle Leiste und vor ihr etwa
fünf Zellen (Abb. 50)............ Ascotis Hübner
- Frontal des Spiraculums am 5. Abdominal-
segment Leisten und Punktgrübchen (Abb.
65, 69), oder nur Leisten (Abb. 68) oder
Punktgrübehen (Ab PINE 40
0 (39) 9. Abdominalsegment mit Punktgrübchen
(Abb. 56,6 Wis. ee 41
- 9. Abdominalsegment ohne Punktgrübchen
(Abb4 62563) ARE 43
41 (40) Frontal des Spiraculums am 5. Abdominal-
segment keine glatte Flache, nur Punktgriib-
Chen (Ab ET) me Chiasmia Hübner
= Frontal bzw. frontodorsal des Spiraculums
am 5. Abdominalsegment eine glatte Fläche
und davor liegen Punktgriibchen bzw. eine
@uerleiste (Abb) 42
42 (41) An der Basis des 5. Abdominalsegmentes la-
teral nur Punktgrübchen ohne eine dunkle
Querleiste (Abb. 59). Stiel am Kremaster re-
lativakurz Abb 58). Parectropis Sato
= An der Basis des 5. Abdominalsegmentes la-
teral eine dunkle Querleiste mit einer Reihe
von dicht gedrangten Punktgriibchen (Abb.
60). Stiel des Kremasters lang (Abb. 61) … …
en ette a ER, Tephrina Gueneé
43 (40) Dorsalrinne ohne Kaudalausläufer … … … … …
br HS. a A Seta OR RES Lo Euconista Lederer
— Dorsalrinne mit Kaudalausläufern … … … 44
44 (43) Frontal des Spiraculums am 5. Abdominal-
segment liegen nur Punktgrübchen (Abb.
= Frontal des Spiraculum am 5. Abdominalseg-
ment liegt eine glatte Flache, dann Punkt-
erubcheny(Abbyp 9) E O 46
— Frontal des Spiraculums am 5. Abdominal-
segment liegt zuerst eine glatte Fläche, dann
eine dicke Leiste und eine Vertiefung mit
Leisten und Punktgrübchen (Abb. 226).
a onl En Odontognophos Wehrli
45 (44) Kaudal des thorakalen Spiraculums eine
längliche tomentöse Erhebung (vgl. Abb. 31)
aA et De TS de ee Isturgia Hübner
Abb. 58-87. — 58, 59, Parectropis similaria; 60, 61 Tephrina arenacearia; 62, Deileptenia ribeata; 63, Arichanna melanaria; 64,
Stegania dilectaria; 65, Peribatodes rhomboidaria; 66, Semiothisa signaria; 67, 68, Itame brunneata; 69, 71, Diastictis artesiaria;
70, Narraga fasciolaria, 72, Abraxas grossulariata, 73, Ennomos autumnaria, 74, 76, Apeira syringaria, 75, 77, 78, Hemistola
biliosata; 79, Jodis putata; 80, 81, Pseudoterpna pruinata; 82-84, Thalera fimbrialis, 85, Chlorissa cloraria, 86, Hemithea aesti-
varia, 87, Epirrita christyi.
58, 61-63, 66, 67, 70-73, 80, Abdominalende in Dorsalsicht;
59, 60, 64, 65, 68, 69, 75, 78, 89, Basis des 5.
Abdominalsegmentes in Lateralsicht; 75, 79, Abdominalende in Ventralsicht; 76, 82, 86, Habitusbild in Ventralsicht; 77,
Kopf und Beine in Ventralsicht; 81, Abdominalende in Lateralsicht; 84, 85, Augen und Umgebung.
52)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
36
— Kaudal des thorakalen Spiraculums keine to-
mentöse Erhebung, nur grobe Runzeln… …
En NEST ES EMCI Lobophora Curtis
46 (44) Dorsalrinne mit 8-9 mittelgroßen kaudalen
Ausbuchtungen (Abb. 62). Lateraleinschnitt
ohne Runzeln um die Spitze. Ventralseite des
Kremasters mit großen Basalhöckern ............
nt de Deileptenia Hübner
= Dorsalrinne mit etwa 20 kleinen Ausbuch-
tungen (Abb. 63). Lateraleinschnitt mit star-
ken Runzeln um die Spitze. Basalhöcker an
der Ventralseite des Kremasters schwach oder
fehlenden pers Arichanna Moore
47 (32) 5. Abdominalsegment frontal des Spiracu-
lums mit einer schlank ovalen Leiste. Auch
der Kaudalrand des 4. Segmentes dort ver-
diekaAbbrCA)E mel nn 48
- 5. Abdominalsegment frontal des Spiracu-
lums mit etwa fünf Zellen, die kaudal durch
eine dicke Querleiste begrenzt sind (Abb. 50)
Dea Centa SONICS Ei REA Cleora Curtis
- 5. Abdominalsegment frontal des Spiracu-
lums mit Punktgriibchen und/oder Leisten
(ABS AGO ICS ACSAGO) ren... 49
48 (47) Puppe 6-8 X 2,5 mm........ Stegania Gueneé
= Ruppe 92103 MEI Lygdia Gueneé
49 (47) Frontal des Spiraculums am 5. Abdominal-
segment eine fast skulpturlose, glatte Flache
und vor ihr Punktgrübchen bzw. Leisten
ADD IAG). RI AUR os 50
— Frontal des Spiraculums am 5. Abdominal-
segment Punktgrübchen und/oder Leisten,
ohne eine glatte Fläche dazwischen (Abb. 57,
GSCI) See na A EEE ns 52
50 (49) 9. Abdominalsegment mit Punktgrübchen
(GElPABEAGID) Re nn an! Alcis Curtis
— 9. Abdominalsegment ohne Punktgrübchen
(Vel Abba ennio a er 51
51 (50) Lateral an der Basis des 5. Abdominalseg-
mentes eine dicke Querleiste und vor ihr
dichte Punktgrübchen (Abb. 65) … … … … …
MERE RACE RATS. RNL CD, Peribatodes Wehrli
= Lateral an der Basis des 5. Abdominalseg-
mentes nur Punktgrübchen, keine Querleiste
(ADI) tT Ik Paradarsia Warren
52 (49) An der Basis des 5. Abdominalsegmentes la-
teral nur Punktgriibchen (Abb. 57) … … … 53
Patocka: Puppen der Spanner Mitteleuropas
= An der Basis des 5. Abdominalsegmentes la-
teral Leisten und dazwischen manchmal
Punktgrübchen (Abb. 68, 69).................. 54
53 (52) Stiel des Kremasters lang (Abb. 66). Ventral-
seite des Kremasters ohne deutliche Basal-
hécker. Metanotum ohne Punktgrübchen
(velFAbb SO) Semiothisa Hübner
— Stiel des Kremasters kurz (Abb. 70). Ventral-
seite des Kremasters mit deutlichen Basal-
höckern. Metanotum mit Punktgrübchen
(vel MADD 2 TD) ER Narraga Walker
54 (52) Stiel des Kremasters lang und schlank (Abb.
VORO RR Ne Diastictis Hübner
— Stiel des Kremasters kurz und dick (Abb. 67)
PA E DRS AE Se EERE Itame Hübner
Puppe glänzend braunschwarz, Kaudalstrei-
fen des 5.-8. Abdominalsegmentes leuchtend
gelb. Häkchen am Kremaster gleich groß,
dick, zerstreut, Sd, entspringt nahe der Basis
des Kremasters (Abb. 72)....... Abraxas Leach
— Puppe nicht glänzend schwärzlich mit gelben
2)
Seitenstreifen. Wenn die Häkchen am
Kremaster gleich groß und dick sind, ent-
springt Sd, nicht nahe der Basis des
Riremasters en eee RO 56
56 (55) Mittelteil der Puppe stark verbreitert (Abb.
76) Thorax und Abdomen dorsal mit dichten
winzigen Börstchen besetzt (Abb. 74) … … …
dente RN PIGRI RIE LIRE PONI Apeira Gist
= Puppe nicht verbreitert, ohne Börstchen … …
REY À IERI. NOE AMR A RO 57
57 (56) Am 9. Abdominalsegment an Stelle der
Borsten D, (und manchmal auch D,) zahnar-
tige Spitzen. 9. Abdominalsegment dorsal
stark langsgefurcht (Abb. 73). Puppen etwa
15-22 mm lang, matt.... Ennomos Treitschke
— Am 9. Abdominalsegment D' borstenartig
oder rückgebildet. Puppe oft kleiner und/-
oderolinzendi nee 58
58 (57) Kremaster ventral stark radial gefurcht, und/-
oder seine Borsten, auch D,, entspringen fä-
cherartig von den Seiten der Spitze, sind +
gleich groß (D, manchmal etwas stärker) und
am Ende stark spiralig eingerollt (Abb. 75,
79, 80). Vorderschenkel nicht sichtbar oder
klein (Abb. 82, 86, 88). Puppen bunt, oft
grün gefärbt und/oder mit Zeichnungen.......
Abb. 88-114. — 88, Chlorissa cloraria; 89, Epirrita christyi; 90, Larentia clavaria, 91, Selenia dentaria; 92, Hypoxystis pluviaria;
93, Plagodis pulveraria, 94, Opisthograptis luteolata, 94a, Therapis flavicaria, 95, 100, Angerona prunaria, 96, 101, Crocallis
elinguaria, 97, Epione parallelaria, 98, Synopsia sociaria, 99, Aspilates gilvaria, 102, Epione repandaria; 103, 105-107, Dyscia
conspersaria; 104, 108, Pseudopanthera macularia; 109, Petrophora chlorosata; 110-112, Epirrhanthis diversata; 113, Hylaea fa-
sciaria, 114, Odontopera bidentata.
88, 94, 94a, Habitusbild in Ventralsicht; 89, 90, 105, 108, 109, 110, 114, Abdominalende in Dorsalsicht; 91, 106, 111, 113,
Erhebung kaudal von dem thorakalen Spiraculum; 92, Basis des 5. Abdominalsegmentes in Lateralsicht; 93, 107,
Abdominalende in Lateralsicht; 95, 96, Augen und Umgebung; 97, Lateraleinschnitt und Dorsalrinne in Lateralsicht; 98-
101, Kopf und Beine in Ventralsicht; 102-104, Metanotum, 1. Abdominalsegment; 112, Labrum, Labium.
DI
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
PN EI ah Route one AE occ 59
= Kremaster anders gestaltet, ventral meist
nicht regelmäßig radial gefurcht. D, wenn
vorhanden, entspringt von seiner Dorsalseite.
Borsten oft ungleich groß oder lang. Im
Zweifelsfall Vorderschenkel groß (Abb. 1),
Puppen oft zeichnungslos und + glänzend .....
59 (58) Proboscis so lang wie die Mittelbeine (Abb.
82, 86). Abdomen ohne stachelige Skulptur.
RAN RE IAA TNT, 63
= Proboscis kiirzer als die Mittelbeine, doch
länger als die Hälfte dieser (Abb. 77).
Wenigstens die mittleren Abdominalsegmen-
te mit stacheliger Skulptur (Abb. 78) … … 60
= Proboscis kürzer als die Hälfte der Mittel-
beine. Abdominalsegmente ohne stachelige
Skulpturs a0. ee ane Aplasta Hübner
60 (59) Puppe einfarbig grün, Exuvie weißlich grün-
CEE sia Hemistola Warren
= Puppe mit Zeichnungen und dunklen Pinna-
CUITE MB OISE NE Rene 61
61 (60) Puppe grün, am Rücken z. T. rotbraun, 15-
20 mm lang. Hinterbeine meist nicht sicht-
DAT a Geometra Linnaeus
= Puppe nicht griin gefärbt, 11-14 mm lang.
Spitze der Hinterbeine sichtbar, groß ...... 62
62 (61) Fliigeladerung deutlich schwarz abgesetzt,
Grundfarbung weißgräulich........................
Et aad Thetidia Boisduval
— Flügeladern kaum dunkler, Grundfärbung
hell rostbraun. ................ Comibaena Hübner
63 (59) Kremaster mittellang mit 4 Paaren von Häk-
chen (Abbi80) pee: er re au 64
— Kremaster sehr lang mit 5-7 Paaren von
Häkchen (Abb. 79) ................. Jodis Hübner
64 (63) Dorsalrinne und Lateraleinschnitt vorhan-
den, deutlich (Abb. 80, 81) ..........................
EERE N. Pseudoterpna Hübner
= Dorsalrinne und Lateraleinschnitt nicht vor-
handen (vel ADD A95) pee eee eee 65
65 (64) Puppe recht schlank, mehr als 4 mal so lang
wie breit. Frontalhöcker relativ groß (Abb.
86). Puppe eintönig sandfarben oder grün-
lich, nur der Rückenstreifen am 1.-8.
Abdominalsegment dunkel. Spiracula hell ....
en ee Wer Hemithea Duponchel
= Puppe + gedrungen, Frontalhöcker klein
(Abb. 82, 88). Puppe meist bunter gezeich-
net, auch am Kopf und Thorax. Spiracula
dunkel gefleckt (Abb. 83) ........................ 66
66 (65) Grenze zwischen Vorderbein und Antenne
etwa 2X so lang wie die Grenze zwischen
Vorderbein und Gena (Abb. 84). Kopf und
Thorax ventral, auch die Adern der Vorder-
flügel sehr kontrastreich dunkel gezeichnet.
38
Puppe relativ schlanker, 13-15 mm lang.......
een ee Thalera Hübner
= Grenze zwischen Vorderbein und Antenne
etwa 3X so lang wie die Grenze zwischen
Vorderbein und Gena (Abb. 85). Kopf und
Thorax ventral oft weniger kontrastreich ge-
zeichnet. Puppe mehr gedrungen, 9-11 mm
Riga ni PA dal. Chlorissa Stephens
67 (58) Vorderschenkel nicht sichtbar (Abb. 99, 101)
pat O RO A M ME 68
— Vorderschenkel sichtbar (Abb. 1, 165, 166)
NE EN 98
68 (67) D, am Kremaster + parallel, Spitze hakenför-
mig, wenn divergierend, dann erst im
Kaudalteil (Abb. 109) oder Kremaster kurz
und stumpf abgerundet (Abb. 119) ......... 72
= D, am Kremaster stark divergierend, Spitze
oft mehr dorn- als hakenförmig, manchmal +
gestielt. Kremaster länger und/oder spitz
(Abb. 89, 167, 168, 174)... 69
69 (68) Mittlerer Kaudalausläufer der Dorsalrinne
vergrößert, tief (Abb. 89, 174) ................. 70
= Mittlerer Kaudalausläufer der Dorsalrinne
klein oder flach (Abb. 167, 168, 171) bzw.
Dorsalrinne nicht entwickelt (Abb. 93) ... 73
70 (69) Labium nicht sichtbar (Abb. 87). Kremaster
lang vorgezogen, D, einen Winkel von fast
180° bildend (Abb. 89) ...... Epirrita Hiibner
— Labium grof (vgl. Abb. 135), Stiel des
Kremasters kürzer, D, einen spitzen Winkel
bildendii ani tetano si al
71 (70) Sd, vorhanden, entspringt nahe der Basis des
Kremasters, Kaudalausläufer der Dorsalrinne
Spiez Abb Or Larentia Treitschke
- Sd, am Kremaster fehlt. Kaudalausläufer der
Dorsalrinne stumpf abgerundet (Abb. 174)
dte KERS 131
72 (68) Proboscis überragt deutlich den Kaudalrand
des 4. Abdominalsegmentes (Abb. 94 a) … …
Litri i dee Therapis Hübner
— Proboscis überragt den Kaudalrand des 4.
Abdominalsegmentes nicht (Abb. 94) … 73
73 (72) Dorsalrinne nicht entwickelt (Abb. 93) … 74
— Dorsalrinne entwickelt, mindestens stärker
und tiefer als der Dorsaleinschnitt zwischen
dem 8. und 9. Abdominalsegment (Abb.
1058 LOOM iO) ea eee Eee ese 75
74 (73) Punktgrübchen am Abdomen fehlen (Abb.
92). Proboscis kiirzer als die Vorderbeine
(Gye Nolo GE) eee Hypoxystis Prout
= Punktgriibchen am Abdomen vorhanden
(vgl. Abb. 102). Proboscis länger als die Vor-
derbeine (vgl. Abb. 94) … … Plagodis Hübner
75 (73) Kremaster mit 4 Paaren von Häkchen (Abb.
105, 114). D, + größer als die übrigen Bor-
SENTORI 76
— Kremaster mit 2 Paaren von Häkchen. D,
lang, schlank. Puppe 10-11 mm lang … … ….
soan dec nc, Pachycnemia Stephens
= Kremaster mit 2 Paaren von Häkchen. D,
kürzer, relativ dick (Abb. 164). Puppe 16-18
mlam eee Messe sc Senses SR Colotois Hiibner
76 (75) Adern der Vorderflügel stark hervortretend.
Vorderschenkel meist etwas sichtbar (Abb.
Vi) LUPI ENIT
MR IT Opisthograptis Hübner
— Adern der Vorderflügel nicht oder kaum her-
VOREN ede een REA Vu
77 (76) Vorderbeine grenzen aneinander (Abb. 98,
TO) ee BH ET ER esa 78
— Vorderbeine grenzen nicht aneinander (Abb.
DA, NO) ae E A EA 7
78 (77) Mittelbeine grenzen aneinander (Abb. 98).
Kremaster wenig länger als basal breit...........
ee Synopsia Hübner
= Mittelbeine grenzen nicht aneinander (Abb.
99). Kremaster länger als basal breit (Abb.
IONE eee ats Aspilates Treitschke
79 (77) Mittelbeine grenzen aneinander (Abb. 100,
TON eeste RE I CA SAM 80
= Mittelbeine grenzen nicht aneinander (vgl.
IN ol SVN) PRE OMR Repeat ee et net re 82
80 (79) Kremaster in Lateralsicht stumpf. Dorsalrin-
ne mit flachen, breiten Ausläufern. Puppe
unten Srmmilan ossen. Cleorodes Warren
= Kremaster in Lateralsicht eher spitz. Dorsal-
rinne mit tieferen Kaudalausläufern. Puppe
berms mim lance nee. 81
81 (80) Puppe fein skulpturiert, am Riicken glän-
zend, Augen glatt (Abb. 95). Antennen und
Mittelbeine kaudal stumpf, abgerundet
(AL DLO O) ere Angerona Duponchel
= Puppe grob skulpturiert, am Rücken kaum
glänzend. Augen dorsal grob skulpturiert
(Abb. 96). Antennen und Mittelbeine kaudal
IZ (AMORE
EO Sa Ae Crocallis Treitschke
82 (79) Kremaster (in Dorsalsicht, ohne Häkchen)
fast zweimal so lang als basal breit (Abb. 105,
107). Kaudal der thorakalen Spiracula je eine
relativ große tomentöse Erhebung (Abb. 91,
LOC) ER MR LRU 83
= Kremaster kaum länger oder kürzer als basal
breit. Im Zweifelsfall der Höcker kaudal des
thorakalen Spiraculum klein, unauffällig,
kattmstomentos se 85
83 (82) Dorsalrinne mit starken, abgerundeten Kau-
dalansläuterulvel- ADD AE)".
D ce STE EE Charaspilates Wehrli
= Dorsalrinne mit undeutlichen, kleinen bzw.
mehr spitzen Kaudalausläufern (Abb. 105) ...
win er NER. NER Lui 84
Patoëka: Puppen der Spanner Mitteleuropas
84 (83) 1. Abdominalsegment ohne Punktgrübchen
(Abb. 103). Lateraleinschnitt groß, tief (Abb.
LORI Nese aa Dyscia Hübner
= 1. Abdominalsegment mit starken Punkt-
grübchen (Abb. 102). Lateraleinschnitt
schwach entwickelt oder klein (Abb. 97) …
Re eres MONA RE aaa deu Epione Duponchel
85 (82) Metanotum mit Punktgrübchen (Abb. 104)
= Metanotum ohne Punktgriibchen (Abb. 102,
JOSÉ RES A E RR essen 87
86 (85) Pro- und Mesonotum mit Punktgrübchen
(Abb. 104). D, am Kremaster hakenförmig
(Abb. 108) .............. Pseudopanthera Hübner
= Pro- und Mesonotum ohne Punktgriibchen
(vgl. Abb. 210). D, am Kremaster schlank
dornförmig, in der kaudalen Hälfte vonein-
anderidiversierend (Ab bal 09) Pr ee
ee ce Petrophora Hiibner
87 (85) Thorakale Spiracula kaudal mit je einem gro-
ßen, elliptischen, tomentösen Höcker (Abb.
U) a) ale ii ia al mis Selenia Hiibner
— Thorakale Spiracula kaudal ohne oder nur
mit kleinen und schmalen, bzw. nicht to-
mentösen Höckern (Abb. 111, 113) … … 88
88 (87) 1-2. Abdominalsegment ohne Punktgrüb-
chen (vgl. Abb. 103). D, am Kremaster meist
+ weit voneinander entfernt (Abb. 120-122)
= 1-2. Abdominalsegment mit Punktgriibchen
(vgl. Abb. 102). D,am Kremaster entsprin-
gen nahe beieinander (Abb. 110, 114, 119)
IA di se ns Mes Madsen le ML ie ne al lu qe 89
89 (88) D, am Kremaster in Dorsalsicht deutlich län-
ger als der Kremaster selbst, parallel (Abb.
110). Labrum auffallend lang , abgerundet
(CAB DMC) eee. Epirrhanthis Hübner
= D, am Kremaster nicht länger als der Kre-
master selbst oder distal stark divergierend
(Abb. 114, 115, 119). Labrum meist kurz
undibreie (Ab bela G Ne) eee eee 90
90 (89) 10. Abdominalsegment (ohne Kremaster) in
Dorsalsicht fast 2 mal länger als das 9. Abdo-
minalsegment, mit einer gebogenen, feinen
Querfurche (Abb. 114). Puppe 17-20 mm
EE nn Odontopera Stephens
= 10. Abdominalsegment (ohne Kremaster)
nicht oder wenig länger als das 9. Abdomi-
nalsegment, ohne durchgehende Querfurche.
Puppenjote kurzemalsy ls) mm mes 91
91 (90) Lateraleinschnitt groß, wenigstens bis zur
Mitte des 10. Abdominalsegmentes reichend
(Abb): BEA dele er 92
= Lateraleinschnitt klein und kurz (Abb. 118)
te Di eb On 95
92 (91) Kaudal des thorakalen Spiraculums je eine
39
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
40
längliche, stark tomentöse Erhebung (Abb.
113). Lateraleinschnitt distal schmal. Kre-
master auch im distalen Bereich dorsal skulp-
turiert, Sd, entspringen etwa in seiner Mitte
EE Hylaea Hübner
- Kaudal des thorakalen Spiraculums ebenfalls
je eine längliche tomentöse Erhebung. Late-
raleinschnitt distal + breit. Kremaster dorsal
auch im distalen Bereich skulpturiert, Sd,
entsprinseinahe seinen Basis au...
BEE EN tre eek Menophra Moore
= Kaudal des thorakalen Spiraculum liegt keine
oder eine ganz unauffällige und kaum tomen-
töse Erhebung (Abb. 111). Kremaster im di-
stalen Bereich + glatt. Sd, entspringt meist
nicht so nahe der Kremasterbasis ............. 93
93 (92) Labrum mit wenig konvergierenden Seiten
(Abb. 117). Puppen fein skulpturiert, relativ
stark glänzend. Kremaster im Basalteil
schwach gerunzelt ... Puengeleria Rougemont
= Labrum trapezformig mit stark konvergieren-
den Seiten (Abb. 116). Puppe nur mäfig
glänzend, + gröber skulpturiert. Kremaster
dorsal an der Basis stark gerunzelt… … … 94
94 (93) Puppe 12-20 mm lang. 8. Abdominalseg-
mentionne PUNKEEEUDERENS nee
Te cate diga Campaea Lamarck
- Puppe 8-11 mm lang. 8. Abdominalsegment
mi@starken Punktsrübchen® ee...
BR LE le Cepphis Hübner
95 (91) Sd, entspringen nahe der Basis des Kremas-
ters, dieser kurz, stumpf, abgerundet, D,
groß Abb. 19) #20 Lomographa Hübner
- Sd, entspringen etwa von der Mitte des Kre-
masters, dieser relativ lang, spitz (Abb. 118)
Ri e cio Cabera Treitschke
96 (88) Sd, entspringen am Kremaster auf der Höhe
von D, in der Mitte des Kremasters oder di-
stal davon (Abb. 121, 122). Lateraleinschnitt
SCA len th a e 97
- Sd, entspringen am Kremaster mehr frontal
als D,, nahe der Basis (Abb. 120). Lateralein-
schnitt kurz, breit, stumpf abgerundet ..........
RT à A Mi Sciadia Hübner
D, mehr als halb so lang wie der
Kremaster, dieser kürzer als basal breit.
97 (96)
Patoëka: Puppen der Spanner Mitteleuropas
Dorsalrinne mit 2-3 Kaudalausläufern
(Abb. 224). Labium sehr groß (Abb. 225)
Perconia Hübner
= D, mehr als halb so lang wie der
Kremaster, dieser kiirzer als basal breit.
Dorsalrinne mit mehreren Kaudalausläu-
fern. (Abb-122)MEabiumierote ne
Le RU Gnophos Treitschke s. lat.
— D, weniger als halb so lang wie der Kre-
master, dieser langer als basal breit. Dorsal-
rinne mit mehreren Kaudalausläufern
(Abb. 121). Labium groß Crocota Hübner
An der Basis der Vorderflügel je ein starker
Hocker (Abbal23 5125.30) eee 99
= An der Basis der Vorderflügel kein Höcker
CAD 82) ee Somer SNR 101
Proboscis überragt den Kaudalrand des 4.
Abdominalsegmentes (Abb. 123).
Kremaster mit relativ langen Borsten (Abb.
DA) er tee eae Rhodometra Meyrick
— Proboscis überragt den Kaudalrand des 4.
Abdominalsegmentes nicht (Abb. 125,
130). Kremaster mit kurzen Borsten (Abb.
WORSE E Tee mn 100
Frons mit langem, zweispitzigem Fortsatz
(Abb. 125). Die Borsten am Ende des
Kremasters einzeln (Abb. 127) …
liel Timandra Duponchel
— Frons ohne Fortsatz (Abb. 130). Die
Borsten am Ende des Kremasters in einer
Gruppe (Abb. 128) .... Cyclophora Hiibner
Puppe + matt, griin oder leder- bzw. sand-
farben, dann mit dunkler Sprenkelung
oder anderer Zeichnung bzw. schwarzen
Spiracula. Exuvie grünweiß, gelbweiß oder
trüb;bräunlichgelbe a eee 102
= Puppe + glänzend, gelb-, rot- bis schwarz-
Braun, zeichnungslos rer 113
102 (101) Frontolaterale Lappen des Metanotums
spitz Abb. 153)... nme. ee 128
— Frontolaterale Lappen des Metanotums
abgerundet (Abby 195) ee ee: 165
103 (165) Adern der Vorderflügel stark hervortre-
tend RO QUE A SE dl Oi n I 76
= Adern der Vorderfliigel nicht stark hervor-
tretend 0 LO eee 104
100 (99)
101 (98)
Abb. 115-145. — 115, 117, Puengeleria capreolaria; 116, Campaea honoraria; 118, Cabera pusaria; 119, Lomographa temera-
ta, 120, Sciadia tenebraria, 121, Crocota nivearia, 122, Gnophos dilucidaria, 123, 124, Rhodometra sacraria; 125-127,
Timandra griseata, 128, 130, Cyclophora linearia, 129, Ourapteryx sambucaria, 131, Eulithis populata, 132, 133, Ecliptopera
silaceata; 134, Electrophaes corylata; 135, Dysstroma truncata; 136, Cidaria fulvata; 137, Thera variata; 138, Rheumaptera cer-
vinalis, 139, 140, R undulata; 141, Philereme transversata, 142, Triphosa dubitata, 143, Pterapherapteryx sexalata; 144,
Xanthorhoe designata; 145, Epirrhoe alternata.
115-118, 137, Abdominalende in Lateralsicht; 116, 117, 133, 135, Labrum, Labium; 119-122, 124, 127-129, 131, 132,
134, 136, 143, Abdominalende in Dorsalsicht; 123, 125, 130, Habitusbild in Ventralsicht; 126, Kopffortsatz in Lateralsicht
(Umriss); 138, 140, 141, 142, Basis des 5. Abdominalsegmentes in Lateralsicht; 139, Basis des 5. Abdominalsegmentes,
Dorsalsicht; 144, 145, Auge und Umgebung.
4]
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
146
42
o
RS 288 So] 53
104 (103) D, am Kremaster an einem kurzen Stiel,
eine Winkel von fast 90° bildend (Abb.
129) ARuppelibertliimmilang an.
AR a eben OLA Ourapteryx Leach
= D, am Kremaster nicht oder kaum gestielt.
Puppe unter 16 mm lang................... 105
105 (104) Kremaster mit 4 Paaren von Borsten (Abb.
LAS TESA Re NT A SIREN ae ae RE 106
= Kremaster mit 6 Paaren von Borsten (Abb.
ISOMEREN ento 111
106 (105) 10. Abdominalsegment dorsal stark ver-
kürzt und + längsgefurcht oder längsge-
rippt (Abb. 131). Puppen über 12, oft über
IRR Eulithis Hübner
= 10. Abdominalsegment stark verkiirzt, je-
doch nicht längsgefurcht. Puppenlänge
unter 12 mm....... Xanthorhoe Hübner p.p
(X. munitata (Hübner))
= 10. Abdominalsegment nicht stark ver-
kürzt, kaum längsgefurcht (Abb. 132).......
PRR RE ER Se TER 107
107 (106) Labium nicht sichtbar oder ganz klein
(Abb 3 200) een eek 108
— Labium relativ groß (Abb. 135) ......... 110
108 (107) Dorsalrinne ohne Kaudalausläufer (Abb.
152 depend Se Ecliptopera Warren
= Dorsalrinne mit Kaudalausläufern, wenig-
stens der mittlere gut sichtbar (Abb. 134,
DOSE aaa PRE Hamer e 109
109 (108) Dorsalrinne mit deutlichen spitzen tomen-
tösen Ausläufern (Abb. 134). Puppe deut-
lich dunkel gefleckt...... Electrophaes Prout
= Dorsalrinne mit undeutlichen Ausläufern,
nur der mittlere etwas mehr ausgepragt
(Abb. 208). Puppe undeutlich oder nicht
pelleckuse:. een... 156
110 (107) Puppe dünnschalig, nicht gesprenkelt,
nicht bereift, Exuvie matt gelblich ............
gm koala: Dysstroma Hiibner
— Puppe dickschalig, dunkel gesprenkelt,
deutlich bereift, Exuvie hellbraun. .............
ae Een en adres Chloroclysta Hübner
- Puppe dickschalig, braun mit Zeichnun-
gen, insbesondere an den Adern der Vor-
derfliigel, nicht bereift, nur 7-8 mm lang
RES A PR SRI LL SEI ERI nia 150
Patoëka: Puppen der Spanner Mitteleuropas
111 (105) Dorsalrinne und Lateraleinschnitt nicht
sichtbar (Abb. 136)..... Cidaria Treitschke
= Dorsalrinne und Lateraleinschnitt + sicht-
loer (lolo, 197) HEN RR ID
112 (111) Borsten am Kremaster satt braunrot, Exu-
vie weißlichgelb. Dorsalrinne schwach,
meist nur mit einzelnen, kleinen Kaudal-
ausläntern I Plemyria Hübner
— Borsten am Kremaster bräunlich. Exuvie
bräunlich weiß oder gelb. Dorsalrinne mit
mehreren meist deutlichen Kaudalausläu-
fern Abba Se Thera Stephens
113 (101) An der Basis des 5. Abdominalsegmentes,
dorsal oder lateral, dunkle Leisten, die
Querrinnen oder -vertiefungen begrenzen
(Abb.138142 eisten re 114
= An der Basis des 5. Abdominalsegmentes
keine Leisten, Querrinnen oder -vertiefun-
gen, hôchstens dichte Punktgriibchen
CAB yi2 2) pet ai aan 116
114 (113) An der Dorsalseite der Basis des 5.
Abdominalsegmentes Querleisten und da-
zwischen eine Querrinne. Sie laufen lateral
+ in die Nähe der Spiracula herab (Abb.
158-140) ee Rheumaptera Hübner
= Nur lateral an der Basis des 5. Abdominal-
segmentes Querleisten und Vertiefungen
(ABB AA OTA DID EE 115
115 (114) 1-2 laterale Vertiefungen am 5. Abdomi-
nalsegment ungeteilt (Abb 141)................
A A ee ee Philereme Hübner
= Laterale Vertiefung an der Basis des 5. Ab-
dominalsegmentes in mehrere Zellen un-
terteilt (Abb. 142)........ Triphosa Stephens
116 (113) Am Kremaster - außer den Borsten D,, die
starke Endhäkchen, Enddornen bzw. eine
gestielte Endgabel bilden - gibt es nur ein
Paar kleiner Borsten (L, oder D,) (Abb.
WAZ 146, FO) PN 117
- Am Kremaster, außer D, noch 2-3 Paaren
von Borsten (Abb. 167, 183, 201, 218) ....
ee sent PAR et ra Se D ER Di 123
117 (116) Kremaster mit einem Paar von lateralen
Höckern. D, zusammen gestielt, stark di-
vereierend {ADD AS) ES
vs eee ahaha lola Pterapherapteryx Curtis
Abb. 146-178. — 146, Catarhoe cucullata; 147, C. rubidata; 148, Epirrhoe alternata, 149, 150, Nycterosea obstipata, 151, 152,
Camptogramma bilineata; 153, 154, Scopula marginepunctata, 155, Idaea aureolaria, 156, I. rufaria, 157, I. deversaria, 158,
159, Rhodostrophia vibicaria, 160, 161, Baptria tibiale, 162, Glacies alticolaria, 163, Gnophos unicoloraria, 164, Colotois pen-
naria; 165, 167, Nothacasis sertata, 166, 171-173, Anticollix sparsata, 168, Acasis appensata, 169, Trichopteryx polycommata;
170, 7. carpinata, 174-176, Anticles badiata; 177, Discoloxia blomeri, 178, Venusia cambrica.
146, 148, 150, 152, 154, 157, 159, 161-164, 167, 168, 171, 172, 174, 177, 178, Abdominalende in Dorsalsicht; 147,
Dorsalrinne; 149, 151, 153, Pronotum, 1. Abdominalsegment; 158, Erhebung kaudal von dem thorakalen Spiraculum; 160,
Metanotum, 1. Abdominalsegment; 165, Kopf und Beine in Ventralsicht; 166, Vorderschenkel; 169, 170, 175,
Abdominalende in Lateralsicht; 176, Punktgriibchen an der Basis des 2. Abdominalsegmentes.
43
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
= Kremaster ohne Lateralhöcker. Enddornen
schlanker, nicht so lang gestielt, weniger
divergierend (Abb. 146, 150)............. 118
118 (117) Grenzen zwischen Vorderbeine und An-
tennen viel länger als die Grenzen zwi-
schen Vorderbeine und Augen. Grenzen
zwischen Antennen und Augen kaum län-
ger als die zwischen Antennen und Vorder-
beine (ADD Al 44) elena. 119
= Grenzen zwischen Vorderbeine und
Antennen nicht oder wenig länger als die
Grenzen zwischen Vorderbeine und Au-
gen. Grenzen zwischen Antennen und
Augen deutlich langer als die zwischen
Antennen und Vorderbeine (Abb. 145) ....
SOE GATA EEN ENEN Ne ee 120
119 (118) Am Kremaster außer D, noch L, vorhan-
den, nahe bei D, (vgl. Abb. 148) … … … ……
teo AR Xanthorhoe Hübner
— Am Kremaster außer D, noch D,, von D,
enttemti ona nr RIEL 120
120 (118) Dorsalrinne nur mit einem großen Kau-
dal- und Frontalausläufer in der Mitte
(Abb. 146, 147) .......... Catarhoe Herbulot
— Dorsalrinne mit mehreren Kaudalausläu-
fern (Ab bl 485 ONS 2) een. li
121 (120) D, so lang wie oder länger als der Kremas-
ter (Abb. 148)................ Epirrhoe Hübner
- D, kürzer als der Kremaster, dieser + länger
als basal breit (Abb. 150, 152) ........... 122
122 (121) Frontolaterale Lappen des Metanotums
spitz (Abb. 149). Dorsalrinne im Mittelteil
wenig verbreitert (Abb. 150).....................
EE E, Nyeterosea Hulst
- Frontolaterale Lappen des Metanotums
abgerundet. Dorsalrinne im Mittelteil
stark verbreitert (Abb. 152).......................
Se AN. ee Camptogramma Stephens
123 (116) Frontolaterale Lappen des Metanotums
spitzwinkelig (Abb. 153, 160). (Wenn je-
doch das 1. Abdominalsegment mit deutli-
chen Punktgrübchen versehen und die
Skulptur relativ grob ist, suche weiter un-
ten) 124
_ Frontolaterale Lappen des Metanotums
stumpf, abgerundet (Abb. 151, 181).. 129
124 (123) D, gleichlang und selten starker als die
iibrigen Borsten des Kremasters (Abb.
WS pen AED re last 125
= D, deutlich länger als die übrigen Borsten
des Kremasters (Abb. 154, 159, 161) … …
Eene Nile al she 126
125 (124) Dorsalrinne nur mit einem mittelgroßen
Vorsprung in der Mitte. Labium klein
(Abb. 193). Lateraleinschnitt kurz, abge-
zundee.:4.......,. EEE 149
— Dorsalrinne mit kleinen Vorsprüngen, der
mittlere aber sehr lang, bis nahe zur Basis
des Kremasters reichend. (Labium an der
untersuchten Puppe nicht vorhanden) …
lore aloni Glossotrophia Prout
= Dorsalrinne mit mehreren Vorsprüngen,
keiner von ihnen nähert sich der Basis des
Kremasters (Abb. 155, 157). Wenn nur ei-
ner dorsal in der Mitte oder gar keiner vor-
handen ist, dann fehlt das Labium und/-
oder der Lateraleinschnitt sehr groß..........
vi tek si) che nimes SAN pay Idaea Treitschke
126 (124) Kremaster rauh skulpturiert, D, entsprin-
gen weit voneinander entfernt. Vorder-
schenkel klein … … eneen 97
= Kremaster + fein skulpturiert. D, entsprin-
gen nahe beieinander, Vorderschenkel
grol Am 1277
127 (126) Sd, entspringen nahe der Basis des Kre-
masters (Abb. 161). Labium klein (vgl.
ADDII) o Baptria Hübner
= Sd, (wenn vorhanden) entspringen weit
von der Basis des Kremasters entfernt
(Abb. 154, 159). Labium meist groß (vgl.
AA SRE Merle sea 128
128 (127) Kaudal des thorakalen Spiraculums je eine
große, elliptische, stark tomentöse Erhe-
bung (Abb. 158). Puppe sandfarben oder
BRUT nee ice Rhodostrophia Hübner
= Kaudal des thorakalen Spiraculums eine
längliche, weniger auffällige Erhebung
(vgl. Abb. 113). Puppe rotbraun, glänzend
RAN Scopula Schrank
129 (123) Kremaster mit 4 Paaren steifer, gleich gro-
Ser Borsten, am Ende nicht hakenartig. D,
entspringen weit voneinander entfernt
(Abb. 162). Punktgrübchen fehlen............
En TE Glacies Milliere, 1874
= Wenigstens ein Teil der Borsten am
Kremaster hakenartig. Punktgrübchen am
Abdomen + vorhanden … 130
130 (129) Vorderschenkel klein, oft recht schmal.
Labium klein oder fehlend (Abb. 165). D,
am Kremaster länger und stärker als die
übrigen Borsten, + divergierend (Abb.
ZIO Mino 131
= Vorderschenkel, Labium, oder beide mit-
telgroß bis groß (Abb. 1), sonst D, am
Kremaster nicht viel langer oder starker als
die übrigen Borsten. (Abb. 193) … … 135
131 (130) Dorsalrinne in der Mitte beiderseits, insbe-
sondere kaudalwärts, mit abgerundeten
Ausschnitt. Borsten am Enddrittel des
Kremasters konzentriert (Abb. 174).
Punktgrübchen an der Basis des 1.-3. Ab-
dominalsegmentes stark vergrößert (Abb.
(FO) en Anticlea Stephens p.p.
(A. badiata Denis et Schiffermiiller)
= Dorsalrinne mit einem oder mehreren fla-
chen Kaudalen Ausschnitten. Borsten auch
an der Basalhalfte des Kremasters (Abb.
167, 168, 171). Punktgriibchen an der
Basis des 1. -3. Abdominalsegmentes nicht
VER St SA ME ER OCR 132
132 (131) Metanotum mit Punktgrübchen (Abb,
IES) en... 133
= Metanotum ohne Punktgrübchen (Abb.
IES RABEN NE ERE NEE) 134
133 (132)Kremaster in Dorsalsicht länger als basal
breit. Borsten Sd, entspringen nahe der
Basis des Kremasters (Abb. 167) ................
RAR Merete ES Nothacasis Prout
= Kremaster in Dorsalsicht kiirzer als basal
breit. Sd, entspringen etwa in der Mitte
des Kremasters (Abb. 168)...
SS AA RSL LI Acasis Duponchel
134 (132) Puppe ohne oder mit sehr kleinem La-
bium. Lateraleinschnitt undeutlich oder in
der Form eines Griibchens (Abb. 169,
AO ALE LEE Trichopteryx Hübner
= Puppe mit deutlicherem mittelkleinem
Labium. Lateraleinschnitt rinnenförmig,
mit einem kurzen Kaudalausläufer (Abb.
A ahd Ae ETEN Anticollix Prout
135 (130) Kremaster kiirzer als basal breit, sehr grob
skulpturiert (gefurcht, gerippt). D, haken-
formig, kaum divergierend, stärker und
länger als die übrigen Borsten, basal + von-
einander entfernt, der Kremaster dazwi-
schen + konkav (Abb. 163). Epicranial-
naht zwischen Vertex und Frons nicht
sichtbar (Abb. 4). Puppen über 11 mm
= Kremaster kiirzer als breit, stumpf, grob
skulpturiert. D, stärker als die übrigen
Borsten, basal nahe beieinander, ohne Ein-
schnitt dazwischen, ziemlich parallel lau-
fend. Epicranialnaht sichtbar. Puppenlan-
COIN Epilobophora Inoue
= Kremaster oft langer als breit, nur schwach
oder mäßig skulpturiert. D, entspringen
entweder nahe beieinander und divergie-
ren, oder sind nicht größer als die übrigen
Borsten des Kremasters. Puppen manch-
mal auch kiirzer als 10 mm. Epicranialnaht
relativ deutlich (Abb. 3) … … … … … … … 136
136 (135) Kremaster mit 3 Paar Borsten (Abb. 178-
IS OPEN LINE RE 137
= Kremaster mit 4 Paar Borsten (Abb. 190,
DDE IMS IMG) erde deden 144
137 (136) Lateraleinschnitt nicht entwickelt, oder als
einfache Rinne ohne einen größeren
Patocka: Puppen der Spanner Mitteleuropas
Kaudalausläufer (Abb. 175, 182) … … 138
= Lateraleinschnitt deutlich entwickelt, mit
einem starken Kaudalausläufer (Abb. 184)
en Ri ele I ee 142
138 (137) Borsten am Kremaster gleich groß (Abb.
709) ct de Gymnoscelis Mabille
— D, viel größer als die übrigen Borsten am
Kremaster (Abb. 180) … … … ……… … … 139
139 (138) Punktgrübchen an der Basis des 1.-4. Ab-
dominalsegment stark vergrößert und ver-
He te (Abbi YO) Pe 140
— Punktgrübchen an der Basis des 1-4.
Abdominalsegmentes nicht vergrößert
(ADD: 15 TU Er. SIA 141
140 (139) Kremaster am Ende abgerundet, schaufel-
formig (Abb. 177). Puppe 7-8 mm lang....
He NEN EN Discoloxia Warren
= Kremaster am Ende spitz (Abb. 178).
Puppe 8-9 mm lang........... Venusia Curtis
141 (139) Metanotum ohne Punktgrübchen (vgl.
Abb NOA) PER Euchoeca Hübner
— Metanotum mit Punktgriibchen (Abb.
USA) eae SR Hydrelia Hübner
142 (137) Kremaster in Dorsalsicht nicht länger als
basal breit (Abb. 183)... Euphyia Hübner
— Kremaster in Dorsalsicht länger als basal
breit (Abba ISA) ee eee 143
143 (142) Dorsalrinne nur mit drei Kaudalausläufern
(ADD ASS) een Mesoleuca Hübner
— Dorsalrinne mit mehreren Kaudalausläu-
han (Abo, 116) en 146
144 (136) Puppenlänge über 11 mm. Punktgrübchen
fehlen am Metanotum und meist auch am
1. Abdominalsegment (Abb. 187, 219);
falls vorhanden, dann Kremaster recht lang
und die Borsten am 9. Abdominalsegment
deutlich (Abb. 186) … 145
= Metanotum oder wenigstens das 1. Abdo-
minalsegment mit deutlichen Punktgrüb-
chen, oder Puppen unter 10 mm lang.......
145 (144) D, am Kremaster entspringt weit ventral
VON SAN ADR AIG 220) re 146
- D, entspringt etwa auf derselben Höhe wie
SAAB SMS) eee eee 147
146 (145) Kremaster lang zungenförmig (Abb. 186).
Borsten am 9. Abdominalsegment relativ
kräftig (Abb. 186) … Scotopteryx Hiibner
= Kremaster in Dorsalsicht dreieckig, nicht
länger als basal breit (Abb. 220)................
a Coie ane N ER Ae es Pareulype Herbulot
147 (145) Grenzen zwischen Vorderbeinen und An-
tennen viel länger als zwischen Vorderbei-
nen und Augen; Grenzen zwischen Anten-
nen und Augen kaum länger als zwischen
Antennen und Vorderbeinen (Abb. 144) ..
45
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
46
eae A a Xanthorhoe Hübner p. p.
(X montanata (Denis et Schiffermüller),
X. incursata (Hiibner))
= Grenzen zwischen Vorderbeinen und
Antennen nicht oder kaum länger als zwi-
schen Vorderbeinen und Augen; Grenzen
zwischen Antennen und Augen viel länger
als zwischen Antennen und Vorderbeinen
(Ab Daal) ester nues Entephria Hübner
148 (144) Grenzen zwischen Vorderbeinen und
Antennen deutlich länger als zwischen
Vorderbeinen und Augen (etwa gleich lang
wie die Grenzen mit Augen und Genae zu-
sammen). Grenzen zwischen Antennen
und Augen etwa gleich lang wie zwischen
Antennen und Vorderbeine (Abb. 144,
191). Kleine Arten, unter 9, meist unter 8
mm Länge, Skulptur oft grob, D, am
Kremaster hakenförmig, nicht oder nur
wenig divergierend. (Abb. 196-201)
Lateraleinschnitt und Labium meist gut
entwickelt tat... ner 149
= Grenzen zwischen Vorderbeinen und
Antennen nicht oder kaum langer als zwi-
schen Vorderbeinen und Augen; Grenzen
zwischen Antennen und Augen deutlich
länger als zwischen Antennen und Vorder-
beinen (Abb. 145). Wenn die Puppen un-
ter 8 mm lang, fehlt oft das Labium und
der Lateraleinschnitt. D, oft gabelartig di-
vergierend (Abb. 203-205, 220) … … 151
149 (148) Labium sehr klein (Abb. 192). Dorsalrinne
mit einem einzigen, kleinen Kaudalauslau-
fer (Abb. 193). Borsten am Kremaster +
gleich groß, nur im distalen Drittel (Abb.
193). Frontolaterale Lappen des Metano-
tums ziemlich spitz (Abb. 194) .................
ME en IT DO Calliclystis Dietze
= Labium größer (Abb. 1), Dorsalrinne mit
mehreren Ausläufern (Abb. 196, 199,
200), oder D, deutlich gröfser als die übri-
gen Borsten des Kremasters (Abb. 198).
Frontolaterale Lappen des Metanotums +
Alyexmubovalae (Ab 210) nano. 150
150 (149) Metanotum ohne, 1. Abdominalsegment
ohne oder mit schwach angedeuteten
Punktgrübchen (Abb.195). Dorsalrinne
Patoka: Puppen der Spanner Mitteleuropas
mit kleinen Kaudalausläufern, Borsten am
Kremaster fast gleich groß, nur in der di-
stalemsblalfter (Abbi 6) Eeen
N U Chloroclystis Hübner
= Skulptur + grob, 1. Abdominalsegment
und oft auch das Metanotum (bzw. ganze
Thorax dorsal) mit deutlichen Punktgriib-
chen. Sonst meist entweder die Kaudal-
ausläufer der Dorsalrinne grofs (Abb. 201)
oder D, länger als die übrigen Borsten des
Kremasters (Abb. 197, 198, 200) bzw.
Borsten auch in der Basalhalfte des Kre-
NAS TE LS Er Eupithecia Curtis
151 (148) D, am Kremaster deutlich, weit voneinan-
der entfernt, einem stumpfen Winkel bil-
dend (Abb. 190). Metanotum mit Punkt-
grübehen (Vel ADD 2110)
Bae cat Seo Anticlea Stephens p. p.
(Anticlea erivata (Denis et Schiffermüller))
= D, entspringen nahe beieinander (Abb.
DIN PONS) E E I ENO 152
152 (151) D, in Dorsalsicht langer als der Kremaster,
dornartig, divergierend und + lang gestielt.
Oft kleinere Arten, unter 9 mm Länge
(Abb@203 205) Pees Perizoma Hiibner
- D, in Dorsalsicht nicht länger als der
Kremaster, oft hakenartig und + parallel
laufend (Abb. 213, 216, 223), wenn diver-
gierend (Abb. 218, 220), dann oft größere
Artena 153
153 (152) Labium fehlt, oder sehr klein (vgl. Abb.
87, 209). Dorsalrinne und Lateralein-
schnitt + schwach ausgebildet. D, hakenar-
tig, viel stärker als die übrigen Borsten
(Abb. 207, 208, 211). Oft kleinere Arten,
6-Simmilano.. rie 154
— Labium mittelgroR, bis grof (Abb. 1,
135). Dorsalrinne und Lateraleinschnitt
meist deutlich und tief. D, stärker oder
auch gleichstark wie die übrigen Kremas-
terborstent Pes ores een 157
154 (153) Metanotum mit Punktgrübchen (vgl. Abb.
210). Puppe nur etwa 6-7 lang........... 155
— Metanotum ohne Punktgrübchen (vgl.
Abb. 195). Puppe oft länger als 7 mm …
E A e too 156
155 (154) D, am Kremaster viel näher an L, als an Sd,
Abb. 179-211. — 179, Gymnoscelis rufifasciata; 180, 181, Hydrelia flameolaria, 182, Euchoeca nebulata, 183, Eyphyia, 185,
Mesaleuca albicillata, 186, 187, Scotopteryx moeniata; 188, Xanthorhoe montanata; 189, Entephria caesiata; 190 Anticlea deri-
vata; 191-194, Calliclystis v-ata; 195, 196, Chloroclystis rectangulata; 197, Eupithecia absinthiata; 198, E. pulchellata; 199, E.
tenuiata; 200, E. abietaria; 201, E. pusillata; 202, Perizoma alchemillata, 203, P. bifasciata; 204, P. didymata; 205, P. peralle-
lolineata; 206, P. blandiata; 207, P. incultaria; 208, 209, Eustroma reticulata; 210, 211, Asthena albulata.
179, 180, 183, 185, 186, 188-190, 193, 196-208, 211, Abdominalende in Dorsalsicht; 181, 187, 194, 195, 210,
Metanotum, 1. bzw. 2. Abdominalsegment; 182, 184, Abdominalende in Lateralsicht; 191, Augen und Bereich; 192, 209,
Labrum, Labium.
47
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Abb. 212-225. — 212, Minoa murinata; 213, Cosmorhoe ocellata; 214, Nebula salicata, 215, N. nebulata; 216, Colostygia apta-
ta, 217, Hydriomena ruberata; 218, Lampropteryx otregiata, 219-221, Pareulype berberata, 222, Spargania luctuata, 223,
Melanthia procellata, 224, 225, Perconia strigillaria, 226, 227, Odontognophos dumetata.
212-218, 220, 222-224, Abdominalende in Dorsalsicht; 219, Metanotum, 1-2. Abdominalsegment; 221, 226, Basis der 5.
Abdominalsegmentes in Lateralsicht; 225, Labrium, Labium; 227, Höcker kaudal von dem thorakalen Spiraculum.
48
CRESPO PEN (ALEZIO een...
n a Minoa Treitschke
= D, am Kremaster etwa in der Mitte zwi-
schen Sd, und L, entspringend (Abb. 211)
E I PI URANIO, Asthena Hiibner
156 (154) D, am Kremaster entspringen frontal von
Sd (AbbIZ07) Perizoma Hiibner p.p.
St id (P. incultaria Herrich-Schäffer)
= D, am Kremaster entspringen kaudal von
SARA. 208)... Eustroma Hübner
157 (153) Borsten am Kremaster fast gleich lang und
gleich dick, schlank. Sd, entspringt nahe
der Basis des Kremasters (Abb. 213) .........
OR RE Cosmorhoe Hübner
— Borsten am Kremaster nicht gleich lang
und gleich groß (D, deutlich größer, Abb.
216, 218). Wenn doch gleich groß, dann
entspringt Sd, + auf der Höhe von D,, weit
von der Basis des Kremasters entfernt
CDD PM) VS ae enon 158
158 (157) D, entspringt am Kremaster frontal von
Sd, (Abb. 217)......... Hydriomena Hübner
= D, entspringt + auf der Höhe von Sd,
(GN Do LU) een re 159
= D, entspringt kaudal von Sd,. (Abb. 220,
DDD WDD) | NT pate 161
159 (158) D, nicht, oder nur wenig länger als die
übrigen Borsten (Abb. 214, 216) … … 160
= D, deutlich länger und stärker als die übri-
gen Borsten des Kremasters.......................
RE e RA Orthonama Hiibner
160 (159) D, kaum länger, manchmal etwas stärker
als die übrigen Borsten am Kremaster, die-
ser kaudal + stumpf (Abb. 214, 215) … …
Scop RES AIA, Nebula Bryard
— D, deutlich länger und stärker als die übri-
gen Borsten des Kremasters, dieser kaudal
zugespitzt (Abb. 216) … Calostygia Hübner
161 (158) Metanotum und 1. Abdominalsegment
ohne Punktgrübchen (Abb. 219) … … 146
— Metanotum und 1. Abdominalsegment
mit Punktgrübchen (Abb. 210) … … … 162
162 (161) Kremaster in Dorsalsicht länger als basal
breit, D, oft dornartig, divergierend (Abb.
DUS) Ersen raters Lampropteryx Stephens
= Kremaster in Dorsalsicht nicht langer als
basal breit, D, hakenartig (Abb. 220, 223)
OREN ua 163
163 (162) Kremaster in Dorsalsicht kiirzer als basal
breit hakemancie ADD 225)...
N Ei I Melathia Duponchel
— Kremaster in Dorsalsicht nicht oder kaum
kürzer als basal breit, D, mitunter dornar-
de (All DAD) enter EPE 164
164 (163) Dorsalrinne mit deutlichen Kaudalaus-
läufern, auch oft ein Frontalausläufer in
Patotka: Puppen der Spanner Mitteleuropas
der Mitte vorhanden (Abb. 6). D, + ha-
Ken forme Horisme Hübner
— Dorsalrinne mit relativ kleinen Kaudal-
und keinem Frontalausläufer. D, am
Kremaster eher dornartig, gebogen (Abb.
DD) aa Spargania Guéneé, 1857
165 (102) Proboscis kürzer als die Vorderbeine.
Vorderflügel grenzen kurz aneinander.......
ANTES UHR “eae SL NER Lythria Hübner
= Proboscis länger als die Vorderbeine.
Vorderflügel grenzen nicht aneinander......
DISKUSSION
In der vorliegenden Arbeit wurde auf eine Bestim-
mungstabelle der in den derzeitigen Systemen üblichen
Unterfamilien der Geometridae, sowie der Tribus
nach Herbulot (1962-1963) verzichtet. Die bisherigen
Tabellen zur Bestimmung der Unterfamilien von
Khotko (1977), sowie von McGuffin (1967-1981)
kann man für größere oder kleinere Zahl der Gattun-
gen, jedoch nicht für alle erfolgreich benutzen. Die
Tabelle von Khotko ist allzu kompliziert - zur Bestim-
mung von fünf Unterfamilien sind dort mehr als zwei
Textseiten nötig. Die Tabelle von McGuffin ist einfa-
cher, es gibt jedoch bei der Bestimmung mancher
Puppen der mitteleuropäischen Fauna Schwierig-
keiten, besonders was die Unterscheidung der Un-
terfamilien Sterrhinae und Larentiinae betrifft. Noch
schwieriger ist dies bei den Tribus im Sinne von
Herbulot. Nur ftir einen Teil von ihnen gelang es mir
anhand der Puppenmerkmale eine benutzbare
Diagnose aufzustellen und auch dort ist diese nicht für
alle Gattungen nach dem Herbulot'schen System gül-
tig. Unter die von puppenmorphologischer Hinsicht
uneinheitlichen Tribus gehören z. B. die Pseudoter-
pnini (Unterfamilie Geometrinae) und die Lythriini
(Unterfamilie Larentiinae). Mehrere Gattungen soll-
ten anhand der Puppenmorphologie in andere Tribus
gehören als in welche sie von Herbulot gestellt wur-
den, wie z. B. : Calospilos, Lomaspilis, Lygdia und
Stegania (siehe Patoc’ka 1986a), oder Odontognophos,
Cleorodes, Crocota, Synopsia und Menophra (siehe
Patoc’ka 1993).
Die Gattungen des Herbulot’schen Systems ent-
sprechen dagegen in der Mehrzahl auch auf Grund
der Puppenmerkmale sehr gut. Auf wenige Aus-
nahmen soll hier aufmerksam gemacht werden: Die
Art obstipata weicht von vittata stark ab, es wird für
sie deshalb der Gattungsname Nycterosea (im Sinne
von Forster-Wohlfart 1981) benützt. Ebenso wurde
es mit den Gattungen Dysstroma, Chiasmia, Diastictis
und Phigalia gemacht. Außerdem ist auch die
Gattung Xanthorhoe puppenmorphologisch unein-
heitlich. Insbesondere die Art X. munitata entspricht
49
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
viel besser der gattung Eulithis. Die Arten Xanthorhoe
montanata und incursata weichen von den übrigen
stärker ab. Ähnlich ist es auch mit der Gattung
Perizoma, wo besonders die Art incultaria den übri-
gen puppenmorphologisch unähnlich ist. Stark von-
einander abweichende Arten enthält auch die
Gattung Anticlea. Dagegen sind die größeren oder
großen Gattungen Cyclophora, Sterrha, Idaea und
Eupithecia hinsichtlich der Puppenmorphologie rela-
tiv einheitlich gebaut. Die Verhältnisse in der
Gattung Gnophos s. lat. (vgl. Sauter 1990) werden in
einer besonderen Arbeit (Patoëka im Druck) bespro-
chen.
LITERATUR
Forster, W. & T. A. Wohlfahrt, 1981. Die Schmetterlinge
Mitteleuropas 5, Spanner (Geometridae). — Franckh’sche
Verlagshandlung, Stuttgart: 1-311.
Gustafsson, B. (ed), 1987. Catalogus Lepidopterorum
Sueciae. — Riksmuseet, Stockholm: 1-140.
Herbulot, C., 1962-1963. Mise a jour de la liste des
Geometridae de France. — Alexanor, 2: 117-124, 147-
154, 3: 17-24, 85-93.
Khotko, E. I., 1977. A key to the spanpupae (Lepidoptera,
Geometridae). — Academia Scientiarum Beloruss. SSR,
Minsk: 1-80.
Koehler, W., 1937. Beitrag zur Kenntnis der, unter
Streudecke der Kiefernbestände überwinternden
Schmetterlingspuppen. — Institut des recherches des
foréts dominiales Pologne, Traxaux et comptes rendus
Warszawa, Ser. A. Nr. 29: 1-81
Leraut, P. 1980. Liste systématique et synonymique des
Lépidoptères de France, Belgique et Corse. — Alexanor,
Supplément: 1-334.
Ljungdahl, D., 1919. Nagra puppenskrivningar. — Entomo-
logisk Tidskrift 40: 97-119.
McGuffin, W. C., 1967-1981. Guide to the Geometridae of
Canada. — Memoirs of the Entomological Society of
Canada 50: 1-103, 86: 1-159, 101: 1-191, 117: 1-153.
Mosher, E., 1916. A classification of the Lepidoptera based
on characters of the pupa. — Bulletin of the Illinois State
Laboratory of Natural History 12: 1-159.
Nordstrém, F., E. Wahlgren & A. Tulgren, 1941. Svenska
Fjärilar. — Nordisk Familjeboks Förlags Aktiebolag
Stockholm: 1-354.
Patocka, J., 1978. Zur Puppenmorphologie und -Taxono-
mie der Unterfamilie Ennominae insbesondere der Tri-
bus Bistonini (Lepidoptera, Geometridae). — Vestnik C-
eskoslovenské Spoleénosti Zoologické 42: 143-151.
Patocka, J., 1980. Die Raupen und Puppen der Eichen-
schmetterlinge Mitteleuropas. — Monografién zur ange-
wandren Entomologie, Verl. Paul Parey, Hamburg &
Berlin: 1-188.
Patocka, J. 1980-1983. Beitrag zur Puppentaxonomie der
mitteleuropäischen Larentiinae (Lepidoptera, Geometri-
dae). — Biológia, Bratislava, 35: 97-109, 36: 583-593, 37:
559-570, 38: 117-131.
Patocka, J., 1985. Beitrag zur Kenntnis der Puppen der
Unterfamilie Ennominae (Lepidoptera, Geometridae). —
Biológia, Bratislava, 40: 997-1012.
Patocka, J., 1986a. Zur Kenntnis der Puppen der mitteleu-
50
ropäischen Spanner aus der Tribus Abraxini und Semio-
thisini (Lepidoptera, Geometridae). — Biológia, Bratis-
lava, 41: 579-595.
Patocka, J., 1986b. Zur Kenntnis der Puppen der Tribus
Boarmiini (Lepidoptera, Geometridae) in Mitteleuropa.
— Acta Entomologica Bohemoslovaca 83: 301-315.
Patoëka, J., 1992. Uber einige Puppen der Spanner aus der
Tribus Ennomini (Lepidoptera, Geometridae, Ennomi-
nae). — Entomologische Berichten, Amsterdam 52: 171-
176.
Patocka, J., 1993. Uber einige Puppen der Spanner aus der
Tribus Boarmini (Lepidoptera, Geometridae, Enno-
minae). — Entomologische Berichten, Amsterdam 53:
114-120.
Patocka, J., 1994. Die Puppen der mittel- und westeuropäi-
schen Spanner aus der Tribus Asthenini (Lepidoptera,
Geometridae, Larentinae). — Entomologische Berichten,
Amsterdam 54: 13-19.
Patoëka, J., im Druck. Die Puppen der Gnophos-Gruppe
(Lepidoptera, Geometridae) Mitteleuropas. — Zeitschrift
der Arbeitsgemeinschaft österreichischen Entomologen:
im Druck. o
Patocka, J., J. Burgan, M. Capek & M. Stolina, 1960. Die
Tannenschmetterlinge der Slowakei. — Ed. SAV Bratis-
lava: 1-214.
Sauter, W., 1990. Zur Systematik der Gnophos-Gruppe
(Lepidoptera, Geometridae). — Nota Lepidopterologica
12: 328-343.
Speyer, W., 1958. Lepidopteren-Puppen an Obstgewachsen
und in ihrer naheren Umgebung. — Mitteilungen aus der
Biologischen Bundesanstalt fiir Land- und Forstwirt-
schaft 93: 1-40.
Received: 23 April 1993
Revised manuscript accepted: 10 April 1994
ANHANG
Ubersicht der Gattungen und Arten der Familie Geome-
tridae, deren Puppen als Unterlage fiir die Aufstellung dieser
Gattungstabelle untersucht wurden.
Archiearis Hiibner, 1823
parthenias (Linnaeus, 1761) (Slowakei, Holland)
notha (Hiibner, 1803) (Slowakei)
puella (Esper, 1787) (Slowakei)
Alsophila Hiibner, 1825
aescularia ([Denis & Schiffermüller), 1775) (Slowakei)
quadripunctaria (Esper, 1787) (Slowakei)
Aplasta Hiibner, 1823
ononaria (Fuessly, 1783) (Deutschland)
Pseudoterpna Hiibner, 1823
pruinata (Hufnagel, 1767) (Slowakei)
Geometra Linnaeus, 1758
papilionaria (Linnaeus, 1758) (Böhmen)
Comibaena Hiibner, 1823
bajularia ([Denis & Schiffermüller], 1775) (Slowakei)
Thetidia Boisduval, 1840
smaragdaria (Fabricius, 1787) (Slowakei)
Hemithea Duponchel, 1829
aestivaria (Hübner, 1799) (Slowakei, Böhmen)
Chlorissa Stephens, 1831
viridata (Linnaeus, 1758) (Polen)
cloraria (Hübner, 1813) (Slowakei)
etruscaria (Zeller, 1848) (Slowakei)
Thalera Hübner, 1823
fimbrialis (Scopoli, 1763) (Österreich)
Hemistola Warren, 1893
biliosata (Villers, 1789) (Slowakei)
Jodis Hiibner, 1823
lactearia (Linnaeus, 1758) (Slowakei)
putata (Linnaeus, 1758) (Böhmen, Osterreich)
Cyclophora Hübner, 1822
pendularia (Clerck, 1759) (Slowakei, Böhmen)
albiocelaria (Hübner, 1789) (Slowakei)
annulata (Schulze, 1775) (Slowakei)
albipunctata (Hufnagel, 1767) (Slowakei, Böhmen)
puppillaria (Hübner, 1799) (Frankreich)
ruficiliaria (Herrich-Schaffer, 1855) (Slowakei)
porata (Linnaeus, 1767) (Österreich)
quercimontaria (Bastelberger, 1897) (Slowakei)
punctaria (Linnaeus, 1758) (Slowakei)
suppunctaria (Zeller, 1847) (Slowakei)
linearia (Hübner, 1799) (Slowakei)
Timandra Duponchel, 1829
griseata W. Petersen, 1902 (Slowakei)
Scopula Schrank, 1802
immorata (Linnaeus, 1758) (Slowakei)
umbelaria (Hiibner, 1813) (Osterreich)
virgulata (Denis & Schiffermiiller, 175) (Slowakei)
ornata (Scopoli, 1763) (Slowakei)
rubiginata (Hufnagel, 1767) (Slowakei)
marginepunctata (Goeze, 1781) (Slowakei)
corrivalaria (Kretschmar, 1862) (Deutschland)
immutata (Linnaeus, 1758) (Slowakei)
ternata Schrank, 1802 (Slowakei)
Glossotrophia Prout, 1913
asellaria (Millière, 1868) (Italien)
Idaea Treitschke, 1825
rufaria (Hiibner, 1799) (Osterreich)
serpentata (Hufnagel, 1767) (Slowakei)
aureolaria (Denis & Schiffermiiller, 1775) (Osterreich)
extersaria (Herrich-Schäffer, 1848) (Italien)
typicata (Gueneé, 1857) (Italien)
filicata (Hübner, 1799) (Osterreich)
laevigata (Hiibner, 1799) (Deutschland)
obsoletaria (Rambur, 1833) (Italien)
biselata (Hufnagel, 1767) (Slowakei)
calunetaria (Staudinger, 1859) (Schweiz)
inquinata (Scopoli, 1763) (Deutschland)
dilutaria (Hübner, 1799) (Slowakei)
fuscovenosa (Goeze, 1781) (Slowakei)
Patotka: Puppen der Spanner Mitteleuropas
humiliata (Hufnagel, 1767) (Slowakei)
seriata (Schrank, 1802) (Osterreich)
subsericeata (Haworth, 1809) (Italien)
emarginata (Linnaeus, 1758) (Deutschland)
aversata (Linnaeus, 1758) (Slowakei)
degeneraria (Hübner, 1799) (Slowakei)
deversaria (Herrich-Schäffer, 1847) (Slowakei)
Rhodostrophia Hübner, 1823
vibicaria (Clerck, 1759) (Slowakei)
Rhodometra Meyrick, 1892
sacraria (Linnaeus, 1767) (Bosnien)
Lythria Hübner, 1823
purpuraria (Linnaeus, 1758) (Slowakei)
rotaria (Fabricius, 1798) (Russland)
Cataclysme Hübner, 1825
riguata (Hübner, 1813) (Slowakei)
Phibalapteryx Stephens, 1829
virgata (Hufnagel, 1767) (Slowakei)
Scotopteryx Hiibner, 1825
moeniata (Scopoli, 1763) (Osterreich)
coarctaria ([Denis & Schiffermüller], 1775) (Slowakei)
chenopodiata (Linnaeus, 1758) (Slowakei, Dänemark)
luridata (Hufnagel, 1767) (Slowakei)
Larentia Treitschke, 1825
clavaria (Haworth, 1809) (Polen)
Orthonama Hübner, 1825
vittata (Borkhausen, 1794) (Deutschland)
Nycterosea Hulst, 1896 i
obstipata (Fabricius, 1794) (Osterreich)
Xanthorhoe Hiibner, 1825
biriviata (Borkhausen, 1794) (Slowakei)
designata (Hufnagel, 1767) (Slowakei)
munitata (Hiibner, 1809) (Schweiz)
spadicearia (|Denis & Schiffermüller], 1775) (Slowakei)
ferrugata (Clerck, 1759) (Slowakei)
quadrifasciata (Clerck, 1759) (Slowakei)
montanata ([Denis & Schiffermüller), 1775) (Slowakei)
fluctuata (Linnaeus, 1758) (Slowakei)
incursata (Hiibner, 1813) (Schweiz)
Catarhoe Herbulot, 1951
rubidata (|Denis & Schiffermüller], 1775) (Slowakei)
cuculata (Hufnagel, 1767) (Slowakei)
putridata (Herrich-Schaffer, 1852) (Frankreich)
Epirrhoe Hübner, 1825
hastulata (Hübner, 1813) (Slowakei)
pupillata (Thunberg, 1788) (Deutschland)
tristata (Linnaeus, 1758) (Slowakei)
alternata (Müller, 1764) (Slowakei)
rivata (Hübner, 1813) (Dänemark)
molluginata (Hübner, 1813) (Slowakei)
galiata ([Denis & Schiffermüller], 1775) (Slowakei)
Camptogramma Stephens, 1831
51
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
bilineata (Linnaeus, 1758) (Dänemark, Slowakei)
Entephria Hiibner, 1825
cyanata (Hiibner, 1809) (Schweiz)
flavicinctata (Hübner, 1813) (Deutschland)
infidaria (La Harpe, 1852) (Schweiz)
caesiata (Denis & Schiffermiiller, 1775) (Slowakei)
Anticlea Stephens, 1831
badiata ([Denis & Schiffermüller), 1775) (Slowakei)
derivata (|Denis & Schiffermüller], 1775) (Slowakei)
Mesoleuca Hübner, 1825
albicillata (Linnaeus, 1758) (Slowakei)
Pelurga Hiibner, 1825
comitata (Linnaeus, 1758) (Slowakei)
Lampropteryx Stephens, 1831
suffumata ([Denis & Schiffermüller], 1775) (Slowakei)
otregiata (Metcalfe, 1917) (Mähren)
Cosmorhoe Hiibner, 1825
ocellata (Linnaeus, 1758) (Slowakei)
Nebula Bruand, 1846
salicata (Hiibner, 1799) (Slowakei)
tophaceata (Denis & Schiffermiiller, 1775) (Mähren)
nebulata (Treitschke, 1828) (Osterreich)
achromaria (La Harpe, 1852) (Slowakei)
Eulithis Hübner, 1821
prunata (Linnaeus, 1758) (Slowakei)
testata (Linnaeus, 1761) (Slowakei, Böhmen)
populata (Linnaeus, 1758) (Böhmen)
pyropata (Hübner, 1822) (Polen)
pyraliata ([Denis & Schiffermüller), 1775) (Slowakei)
Ecliptopera Warren, 1894
silaceata ([Denis & Schiffermüller], 1775) (Slowakei)
capitata (Herrich-Schäffer, 1839) (Slowakei)
Chloroclysta Hübner, 1825
siterata (Hufnagel, 1767) (Slowakei)
miata (Linnaeus, 1758) (Slowakei, Deutschland)
Dysstroma Hübner, 1825
citrata (Linnaeus, 1761) (Deutschland, Slowakei)
truncata (Hufnagel, 1767) (Slowakei)
infuscata (Tengström, 1869) (Schweden)
Cidaria Treitschke, 1825
fulvata (Forster, 1771) (Slowakei)
Plemyria Hübner, 1825
rubiginata ({Denis & Schiffermüller], 1775) (Slowakei)
Thera Stephens, 1831
obeliscata (Hübner, 1787) (Slowakei)
variata ({Denis & Schiffermüller), 1775) (Slowakei)
britannica (Turner, 1925) (Slowakei)
stragulata (Hübner, 1809) (Slowakei)
cognata (Thunberg, 1792) (Österreich)
juniperata (Linnaeus, 1758) (Slowakei)
cupressata (Geyer, 1831) (Frankreich)
52
Eustroma Hübner, 1825
reticulata (Denis & Schiffermüller, 1775) (Slowakei)
Electrophaes Prout, 1923
corylata (Thunberg, 1792) (Slowakei)
Colostygia Hübner, 1825
aptata (Hübner, 1813) (Osterreich)
austriacaria (Herrich-Schäffer, 1856) (Osterreich)
tempestaria (Herrich-Schäffer, 1856) (Osterreich)
multistrigaria (Haworth, 1809) (Schweiz)
Hydriomena Hübner, 1825
furcata (Thunberg, 1784) (Slowakei)
impluviata ([Denis & Schiffermüller), 1775) (Slowakei)
ruberata (Freyer, 1831) (Osterreich, Deutschland)
Horisme Hiibner, 1825
vitalbata ([Denis & Schiffermiiller], 1775) (Slowakei,
Osterreich)
tersata ([Denis & Schiffermiiller], 1775) (Slowakei,
Deutschland)
aemulata (Hiibner, 1813) (Slowakei)
aquata (Hiibner, 1813) (Deutschland)
corticata (Treitschke, 1835) (Osterreich)
radicaria (La Harpe, 1855) (Deutschland)
Melanthia Duponchel, 1829
procellata ([Denis & Schiffermüller), 1775) (Slowakei)
alaudaria (Freyer, 1846) (Slowakei)
Pareulype Herbulot, 1951
berberata ([Denis & Schiffermiiller], 1775) (Dänemark,
Slowakei)
Spargania Guenee, 1857
luctuata ([Denis & Schiffermiiller], 1775) (Slowakei)
Rheumaptera Hübner, 1822
hastata (Linnaeus, 1758) (Slowakei)
cervinalis (Scopoli, 1763) (Slowakei)
undulata (Linnaeus, 1758) (Slowakei, Osterreich)
Triphosa Stephens, 1829
dubitata (Linnaeus, 1758) (Slowakei)
sabaudiata (Duponchel, 1830) (Osterreich)
Philereme Hübner, 1825
vetulata ([Denis & Schiffermiiller], 1775) (Slowakei)
transversata (Hufnagel, 1767) (Slowakei)
Euphyia Hiibner, 1825
biangulata (Haworth, 1809) (Slowakei)
unangulata (Haworth, 1809) (Böhmen)
frustata (Treitschke, 1828) (Osterreich)
Epirrita Hübner, 1822
dilutata ([Denis & Schiffermüller], 1775) (Slowakei)
christyi (Allen, 1906) (Slowakei)
autumnata (Borkhausen, 1794) (Slowakei)
Operophtera Hübner, 1825
brumata (Linnaeus, 1758) (Slowakei)
fagata (Scharfenberg, 1805) (Slowakei)
Perizoma Hiibner, 1825
affinitata (Stephens, 1831) (Polen)
alchemillata (Linnaeus, 1758) (Slowakei)
hydrata (Treitschke, 1829) (Schweiz)
blandiata ({Denis & Schiffermüller], 1775) (Slowakei)
flavofasciata (Thunberg, 1792) (Slowakei)
didymata (Linnaeus, 1758) (Slowakei, Bòhmen)
obsoletaria (Herrich-Schäffer, 1838) (Osterreich, Schweiz)
sagittata (Fabricius, 1787) (Dänemark, Böhmen)
incultaria (Herrich-Schäffer, 1848) (Osterreich)
parallelolineata (Retzius, 1783) (Osterreich)
Baptria Hübner, 1825
tibiale (Esper, 1791) (Osterreich)
Eupithecia Curtis, 1825
tenuiata (Hübner, 1813) (Slowakei)
inturbata (Hiibner, 1817) (Schweiz, Deutschland)
haworthiata Doubleday, 1856 (Danemark, Deutschland)
immundata (Lienig & Zeller, 1846) (Slowakei, Deutsch-
land)
plumbeolata (Haworth, 1809) (Deutschland)
abietaria (Goeze, 1781) (Deutschland)
analoga Djakonov, 1926 (Deutschland, Polen, Slowakei)
linariata ([Denis & Schiffermüller), 1775) (Deutschland,
Slowakei)
pulchellata Stephens, 1831 (Deutschland)
pyreneata Mabille, 1871 (Slowakei, Deutschland)
laquaearia Herrich-Schäffer, 1848 (Slowakei, Deutschland)
irriguata (Hübner, 1813) (Slowakei, Deutschland)
liguriata Milliere, 1884 (Frankreich)
exiguata (Hübner, 1813) (Deutschland)
insigniata (Hübner, 1790) (Deutschland)
valerianata (Hübner, 1813) (Slowakei, Deutschland)
pygmaeata (Hübner, 1799) (Deutschland)
undata (Freyer, 1840) (Schweiz)
variostrigata Alpheraky, 1878 (Schweiz)
silenata Assman, 1849 (Böhmen, Deutschland)
carpophagata Staudinger, 1871 (Italien)
venosata (Fabricius, 1787) (Slowakei, Deutschland)
schiefereri Bohatsch, 1893 (Italien)
silenicolata Mabille, 1866 (Italien)
alliaria Staudinger, 1870 (Slowakei)
egenaria Herrich-Schäffer, 1848 (Deutschland)
extraversaria Herrich-Schäffer, 1852 (Deutschland)
centaureata (|Denis & Schiffermiiller], 1775) (Slowakei,
Deutschland)
queneeata Miller, 1862 (Österreich)
gratiosata Herrich-Schäffer, 1861 (Frankreich)
thalictrata (Püngeler, 1902) (Schweiz, Italien)
acteata Walderdorf, 1869 (Deutschland)
selinata Herrich-Schäffer, 1861 (Deutschland, Slowakei)
trisignaria Herrich-Schäffer, 1848 (Slowakei, Deutschland)
intricata (Zetterstedt, 1839) (Slowakei, Deutschland)
veratraria Herrich-Schäffer, 1848 (Slowakei, Deutschland)
cauchiata (Duponchel, 1830) (Deutschland)
satyrata (Hiibner, 1813) (Slowakei, Deutschland)
cretaceata Packard, 1874 (Italien)
pernotata Guenée, 1857 (Schweiz)
absinthiata (Clerck, 1759) (Slowakei, Deutschland)
goossensiata Mabille, 1869 (Danemark, Deutschland)
assimilata Doubleday, 1856 (Slowakei, Deutschland)
expallidata Doubleday, 1856 (Deutschland)
vulgata (Haworth, 1809) (Slowakei, Deutschland)
Patoëka: Puppen der Spanner Mitteleuropas
tripunctaria Herrich-Schäffer, 1852 (Slowakei, Deutsch-
land)
denotata (Hübner, 1813) (Slowakei, Deutschland)
subfuscata (Haworth, 1809) (Slowakei, Deutschland)
icterata (Villers, 1789) (Deutschland)
succenturiata (Linnaeus, 1758) (Deutschland)
denticulata (Treitschke, 1828) (Deutschland)
impurata (Hübner, 1813) (Deutschland)
orphnata W. Petersen, 1909 (Slowakei, Deutschland)
subumbrata ([Denis & Schiffermüller), 1775) (Deutsch-
land)
semigraphata Bruand, 1851 (Deutschland)
millefoliata Rössler, 1866 (Deutschland)
santolinata Mabille, 1871 (Frankreich)
simpliciata (Haworth, 1809) (Slowakei, Deutschland)
sinuosaria Eversmann, 1848 (Slowakei, Polen, Deutschland)
druentiata Dietze, 1902 (Frankreich)
distinctaria (Herrich-Schäffer, 1848) (Slowakei, Deutsch-
land)
graphata (Treitschke, 1828) (Schweiz, Italien, Österreich)
pimpinellata (Hübner, 1813) (Deutschland)
pauxillaria Boisduval, 1840 (Deutschland)
gelidata (Möschler, 1860) (Böhmen, Deutschland)
nanata (Hübner, 1813) (Dänemark, Deutschland)
innotata (Hufnagel, 1767) (Deutschland)
fraxinata Grewe, 1863 (Deutschland)
unedonata Mabille, 1868 (Spanien)
virgaureata Doubleday, 1861 (Slowakei, Deutschland)
abbreviata Stephens, 1831 (Deutschland)
dodoneata Guenee, 1857 (Deutschland)
pusillata (Denis & Schiffermüller), 1775) (Slowakei,
Deutschland)
ericeata (Rambur, 1833) (Slowakei)
phoeniceata (Rambur, 1834) (Spanien)
scopariata (Rambur, 1833) (Frankreich)
oxycedrata (Rambur, 1833) (Italien)
lanceata (Hübner, 1825) (Slowakei, Deutschland)
lariciata (Freyer, 1842) (Slowakei, Deutschland)
tantillaria Boisduval, 1840 (Slowakei, Deutschland)
Gymnoscelis Mabille, 1868
rufifasciata (Haworth, 1809) (Slowakei, Deutschland)
Chloroclystis Hübner, 1825
v-ata (Haworth, 1809) (Slowakei, Deutschland)
chloerata (Mabille, 1870) (Slowakei, Deutschland, Däne-
mark)
rectangulata (Linnaeus, 1758) (Slowakei, Deutschland)
debiliata (Hübner, 1817) (Böhmen, Dänemark, Deutsch-
land)
Anticollix Prout, 1938
sparsata (Treitschke, 1828) (Danemark, Deutschland)
Chesias Treitschke, 1825
legatella ([Denis & Schiffermüller), 1775) (Mähren, Schwe-
den)
Carsia
sororiata (Linnaeus, 1758) (Schweden)
Aplocera Stephens, 1827
plagiata (Linnaeus, 1758) (Slowakei)
efformata (Guenee, 1857) (Dänemark)
praeformata (Hiibner, 1826) (Slowakei)
5)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Odezia Boisduval, 1840
atrata (Linnaeus, 1758) (Deutschland)
Schistostege Hiibner, 1825
treitschkei Kovacs, 1957 (Ungarn) L
decussata (Denis & Schiffermüller, 1775) (Osterreich)
Lithostege Hübner, 1825
griseata ([Denis & Schiffermüller), 1775) (nach Khotko,
1977)
Disciloxia Warren, 1895
blomeri (Curtis, 1832) (Slowakei)
Venusia Curtis, 1839
cambrica (Curtis, 1839) (Slowakei)
Euchoeca Hiibner, 1823
nebulata (Scopoli, 1763) (Slowakei)
Asthena Hiibner, 1825
albulata (Hufnagel, 1767) (Slowakei)
anseraria (Herrich-Schäffer, 1855) (Slowakei)
Hydrelia Hübner, 1825
flammeolaria (Hufnagel, 1767) (Slowakei)
Minoa Treitschke, 1825
murinata (Scopoli, 1763) (Slowakei)
Lobophora Curtis, 1825
halterata (Hufnagel, 1767) (Slowakei)
Trichopteryx Hübner, 1825
polycommata ([Denis & Schiffermüller], 1775) (Slowakei)
carpinata (Borkhausen, 1794) (Slowakei)
Epilobophora Inoue, 1943
sabinata (Geyer, 1831) (Schweiz)
Nothocasis Prout, 1937
sertata (Hiibner, 1817) (Slowakei)
Prerapherapteryx Curtis, 1825
sexalata (Retzius, 1783) (Slowakei)
Acasis Duponchel, 1845
appensata (Eversmann, 1842) (Slowakei)
Abraxas Leach, 1815
grossulariata (Linnaeus, 1758) (Slowakei)
Calospilos Hübner, 1825
sylvata (Scopoli, 1763) (Slowakei)
Lomaspilis Hübner, 1825
marginata (Linnaeus, 1758) (Slowakei)
Ligdia Guenée, 1857
adustata ({Denis & Schiffermüller], 1775) (Slowakei)
Stegania Guenée, 1857
dilectaria (Hiibner, 1790) (Slowakei, Frankreich)
Semiothisa Hübner, 1818
54
notata (Linnaeus, 1758) (Slowakei)
alternaria (Hübner, 1809) (Slowakei)
signaria (Hübner, 1809) (Slowakei)
Chiasmia Hübner, 1823
clathrata (Linnaeus, 1758) (Slowakei)
glarearia (Brahm, 1791) (Slowakei)
Diastictis
artesiaria (Denis & Schiffermüller, 1775) (Slowakei)
Isturgia Hübner, 1823
limbaria (Fabricius, 1775) (Slowakei)
Narraga Walker, 1861
fasciolaria (Hufnagel, 1767) (Slowakei)
Itame Hübner, 1823
wauaria (Linnaeus, 1758) (Slowakei)
brunneata (Thunberg, 1784) (Böhmen, Schweden)
Tephrina Guenée, 1845
arenacearia (Denis & Schiffermüller, 1775) (Slowakei)
murinaria (Denis & Schiffermüller, 1775) (Slowakei)
Enconista Lederer, 1853
miniosata (Duponchel, 1829) (Frankreich)
Pygmaena Boisduval, 1840
fusca (Thunberg, 1792) (Schweiz)
Cepphis Hübner, 1823
advenaria (Hübner, 1799) (Polen)
Petrophora Hübner, 1811
chlorosata (Scopoli, 1763) (Slowakei, Böhmen)
Plagodis Hübner, 1823
pulveraria (Linnaeus, 1758) (Slowakei)
dolabraria (Linnaeus, 1767) (Slowakei)
Pachycnemia Stephens, 1829
hippocastanaria (Hübner, 1799) (Holland)
Opisthograptis Hübner, 1823
luteolata (Linnaeus, 1758) (Slowakei)
Epione Duponchel, 1829
repandaria (Hufnagel, 1767) (Slowakei)
paralellaria ({Denis & Schiffermüller), 1775) (Slowakei)
Pseudopanthera Hübner, 1823
macularia (Linnaeus, 1758) (Slowakei)
Hypoxystis Prout, 1915
pluviaria (Fabricius, 1787) (Slowakei)
Therapis Hübner, 1823 u
flavicaria (Denis & Schiffermüller, 1775) (Österreich)
Epirrhanthis Hübner, 1823
diversata (Denis & Schiffermüller, 1775) (Böhmen)
Ennomos Treitschke, 1825
autumnaria (Werneburg, 1859) (Slowakei)
quercinaria (Hufnagel, 1767) (Slowakei)
alniaria (Linnaeus, 1758) (Slowakei, Deutschland)
fuscantaria (Haworth, 1809) (Osterreich, Deutschland)
erosaria ([Denis & Schiffermiiller], 1775) (Slowakei)
quercaria (Hiibner, 1813) (Italien)
Selenia Hiibner, 1823
dentaria (Fabricius, 1775) (Slowakei)
lunularia (Hübner, 1788) (Slowakei, Böhmen)
tetralunaria (Hufnagel, 1767) (Slowakei)
Apeira Gistl, 1848
syringaria (Linnaeus, 1758) (Böhmen, Slowakei)
Artiora Meyrick, 1892
evonymaria (Denis & Schiffermüller, 1775) (Slowakei)
Odontopera Stephens, 1831
bidentata (Clerck, 1759) (Slowakei)
Crocallis Treitschke, 1825
tusciaria (Borkhausen, 1793) (Deutschland)
elinguaria (Linnaeus, 1758) (Slowakei)
Ourapteryx Leach, 1814
sambucaria (Linnaeus, 1758) (Slowakei, Böhmen)
Colotois Hübner, 1823
pennaria (Linnaeus, 1761) (Slowakei)
Angerona Duponchel, 1829
prunaria (Linnaeus, 1758) (Slowakei)
Chondrosoma i
fiduciaria Anker, 1854 (Osterreich)
Apocheima Hübner, 1825
hispidaria ([Denis & Schiffermüller), 1775) (Slowakei)
Phigalia Duponchel, 1829
pilosaria ([Denis & Schiffermiiller], 1775) (Slowakei)
Lycia Hübner, 1825
hirtaria (Clerck, 1759) (Slowakei)
graecaria (Staudinger, 1870) (Bosnien)
isabellae (Harrison, 1914) (Böhmen)
pomonaria (Hübner, 1790) (Slowakei)
zonaria ([Denis & Schiffermüller], 1775) (Slowakei)
Biston Leach, 1815
strataria (Hufnagel, 1867) (Slowakei)
betularia (Linnaeus, 1758) (Slowakei)
Agriopis Hiibner, 1825
leucophaearia ({Denis & Schiffermüller), 1775) (Slowakei)
bajaria ([Denis & Schiffermiiller], 1775) (Slowakei)
aurantiaria (Hiibner, 1799) (Slowakei)
marginaria (Fabricius, 1776) (Slowakei)
Erannis Hübner, 1825
defoliaria (Clerck, 1759) (Slowakei)
Nychiodes Lederer, 1853
obscuraria (De Viller, 1789) (Frankreich)
Patotka: Puppen der Spanner Mitteleuropas
Eurranthis Hübner, 1823
plumistaria (De Viller, 1789) (Frankreich)
Peribatodes Wehrli, 1943
rhomboidaria ([Denis & Schiffermüller], 1775) (Slowakei)
umbraria (Hübner, 1809) (Italien)
secundaria ([Denis & Schiffermüller], 1775) (Slowakei)
Cleora Curtis, 1825
cinctaria ({Denis & Schiffermiiller], 1775) (Slowakei)
Deileptenia Hiibner, 1825
ribeata (Clerck, 1759) (Slowakei)
Alcis Curtis, 1826
repandata (Linnaeus, 1758) (Slowakei)
maculata (Staudinger, 1890) (Slowakei)
Arichanna Moore, 1868
melanaria (Linnaeus, 1758) (Slowakei, Böhmen)
Hypomecis Hübner, 1821
roboraria ([Denis & Schiffermüller), 1775) (Slowakei)
viertlii (Bohatsch, 1883) (Slowakei)
Serraca Moore, 1887
punctinalis (Scopoli, 1763) (Slowakei)
Fagivorina Wehrli, 1943
arenaria (Hufnagel, 1767) (Slowakei)
Ascotis Hiibner, 1825
selenaria (Denis & Schiffermiiller, 1775) (Slowakei)
Ectropis Hiibner, 1825
crepuscularia ([Denis & Schiffermüller], 1775) (Slowakei)
Paradarsia Warren, 1897
consonaria (Hiibner, 1799) (Slowakei)
Parectropis Sato, 1980
similaria (Hufnagel, 1767) (Slowakei)
Aethalura Mc Dunnough, 1920
punctulata ((Denis & Schiffermüller], 1775) (Slowakei)
Ematurga Lederer, 1853
atomaria (Linnaeus, 1758) (Slowakei)
Tephronia Hübner, 1825
sepiaria (Hufnagel, 1767) (Deutschland)
Odontognophos Wehrli, 1951
dumetata (Treitschke, 1827) (Slowakei)
Selidosema Hiibner, 1823
brunnearia (Villers, 1789) (Böhmen)
plumaria ([Denis & Schiffermiiller], 1775) (Slowakei)
Bupalus Leach, 1815
piniaria (Linnaeus, 1758) (Slowakei)
Crocota Hübner, 1823
lutearia (Fabricius, 1794) (Frankreich, Schweiz)
niveata (Scopoli, 1763) (Osterreich)
55
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Menophra Moore, 1887
nycthemeraria (Geyer, 1831) (Deutschland)
Synopsia Hiibner, 1825
sociaria (Hiibner, 1799) (Slowakei)
Cleorodes Warren, 1894
lichenaria (Hufnagel, 1767) (Deutschland)
Gnophos Treitschke, 1825
furvatus ([Denis & Schiffermüller], 1775) (Slowakei)
obfuscatus ([Denis & Schiffermüller], 1775) (Osterreich)
ambiguatus (Duponchel, 1838) (Osterreich)
pullatus ([Denis & Schiffermüller), 1775) (Österreich)
glaucinarius (Hübner, 1799) (Osterreich)
variegatus (Duponchel, 1830) (Österreich, Schweiz)
intermedia Wehrli, 1917 (Slowakei)
dilucidaria (Denis & Schiffermüller}, 1775) (Österreich,
Schweiz)
serotinaria ([Denis & Schiffermiiller], 1775) (Osterreich,
Schweiz)
zelleraria (Freyer, 1836) (Osterreich)
caelibaria (Herrich-Schäfer, 1852) (Osterreich)
operaria (Hiibner, 1813) (Osterreich)
Sciadia Hübner, 1822 fl
tenebraria (Esper, 1806) (Osterreich, Schweiz)
Glacies Millière, 1874 5
alticolaria (Mann, 1853) (Osterreich)
Siona Duponchel, 1829
lineata (Scopoli, 1763) (Slowakei)
Chariaspilates Wehrli, 1953
56
formosaria (Eversmann, 1837) (nach Khotko, 1977)
Aspitates Treitschke, 1825
gilvaria ((Denis & Schiffermüller), 1775) (Slowakei)
Dyscia Hiibner, 1825
conspersaria (Fabricius, 1775) (Slowakei)
Perconia Hübner, 1823
strigillaria (Hübner, 1787) (Mähren)
Cabera Treitschke, 1825
pusaria (Linnaeus, 1758) (Slowakei)
exanthemata (Scopoli, 1763) (Slowakei)
Lomographa Hübner, 1825
bimaculata (Fabricius, 1775) (Slowakei)
temerata ([Denis & Schiffermüller], 1775) (Slowakei)
Campaea Lamarck, 1816
margaritata (Linnaeus, 1767) (Slowakei)
honoraria ([Denis & Schiffermüller), 1775) (Böhmen)
Hylaea Hübner, 1822
fasciaria (Linnaeus, 1758) (Slowakei)
Puengeleria Rougemont, 1903
capreolaria ([Denis & Schiffermiiller], 1775) (Slowakei)
Lignyoptera
thaumastaria Rebel, 1901 (Bosnien)
Theria Hiibner, 1825
rupicapraria ({Denis & Schiffermiiller], 1775) (Slowakei)
primaria (Haworth, 1809) (Deutschland)
JOHN T. POLHEMUS & Dan A. POLHEMUS
University of Colorado Museum, Englewood, U.S.A. & Bishop Museum, Honolulu, U.S.A.
FOUR NEW GENERA OF MICROVELIINAE
(HETEROPTERA) FROM NEW GUINEA
Polhemus, J. T. & D. A. Polhemus, 1994. Four new genera of Microveliinae (Heteroptera)
from New Guinea. — Tijdschrift voor Entomologie 137: 57-74, figs. 1-30. [ISSN 0040-7496].
Published 15 July 1994.
Four new genera of Microveliinae are described from New Guinea. These genera and their con-
stituent species are as follows: Tanyvelia gen. n. containing type species 7. missim sp. n. from
Papua New Guinea; Aegilipsicola gen. n. containing type species A. rapida sp. n. from Papua
New Guinea; Tarsovelia gen. n. containing type species 7. alta sp. n. from Papua New Guinea,
plus 7: arfak sp. n. from Irian Jaya, and 7. dani sp. n. from Irian Jaya; and Neusterinsifer gen.
n. containing type species N. compacta sp. n. from Papua New Guinea, plus N. sepik sp. n. from
Papua New Guinea, N. cyclops sp. n. from Papua New Guinea, N. nabire sp. n. from Irian Jaya,
and N. gladius sp. n. from Irian Jaya. Habitus figures of the above new genera are provided, ac-
companied by illustrations of the male genital structures and distribution maps for all new spe-
cies.
Correspondence: Dr Dan A. Polhemus, Dept. of Natural Sciences, Bishop Museum, P.O. Box
19000-A, Honolulu, HI. 96817, U.S.A.
Key words. — Veliidae; Microveliinae; New Guinea; taxonomy; new genera; new species; keys;
distribution.
The assemblages of Microveliinae occurring on the
islands of the Malay Archipelago exhibit a profound
shift in taxonomic composition as one progresses
from west to east through the region. On the Greater
Sunda Islands, Celebes and the Philippines such fau-
nas are dominated by members of the genera
Pseudovelia Hoberlandt and Microvelia Westwood,
accompanied by several other more ecologically spe-
cialized and less speciose genera. On New Guinea and
surrounding islands, by contrast, Pseudovelia is absent
and there occurs instead a group of previously unde-
scribed endemic genera which fill the typical
Pseudovelia niches. In addition, New Guinea also
supports numerous endemic species of in the genus
Microvelia sensu lato, but as noted by Andersen
(1982) this generic grouping is almost certainly poly-
phyletic. A detailed analysis on a world basis of the
subgroups contained within it will be necessary be-
fore it can be certain that the Papuan and Australian
species presently held in Microvelia are in fact conge-
neric with the Asian forms.
New Guinea thus represents a major center of di-
versification for the Microveliinae, with a fauna de-
rived independently from that occurring in the Asian
tropics. In the present report we describe four new
genera of endemic Papuan Microveliinae, all of which
are easily separable from Microvelia sensu lato on the
basis of distinct apomorphies. The relationships
among these genera are still unresolved, however,
pending the completion of ongoing revisions of the
diverse Australian and New Guinea Microvelia fau-
nas.
Key to genera of New Guinea Microveliinae
1. Middle tarsi subequal in length to middle tibia
(fig. 9); light markings on hemelytra restricted to
basal angles; fore femur of male at least slightly
modified for phoresy … … … … Tarsovelia gen. n.
— Middle tarsi distinctly shorter than middle tibia
(figs. 1, 5, 17); light markings on hemelytra may
be restricted to basal angles, or may occur also on
distal portions; fore femur may or may not be
modified for phoresy (figs. 3, 6, 24-26) ........... 2
2. Claws extremely long; light hemelytral markings
entirely bright greyish to light bluish pruinose;
gula long, rostral cavity demarcated by strongly
Taisedkearin ag e Aegilipsicola gen. n.
— Claws relatively short; light hemelytral markings
not bright pruinose; gula short, rostral cavity not
demarcated by strongly raised carina ............... 3
3. Antennae extremely long (fig. 1); comb on fore
tibia 3/4 the length of the tibia (fig. 3); only mi-
cropterous morph known ....... Tanyvelia gen. n.
— Antennae not extremely long (fig. 17); comb on
fore tibia less than 3/4 length of tibia (except in
Neuterinsifer compacta), if comb on fore tibia 3/4
length of tibia then male proctiger bearing elon-
gate process and female abdominal tergites VII
57
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
and VIII forming an anal plate (see below); only
apterous and macropterous morphs known, latter
with large hemelytral light markings basally, and
often occurring in all closed cells … … … … … … …
4. Female abdominal tergites VII-VIII deflected
ventrad forming an anal plate (fig. 23); male gen-
italia highly modified, bearing a long sinuate an-
teriorly directed sword-like process (figs. 18-22)
aste ee Slee ten. Neusterinsifer gen. n.
— Female abdominal tergites VII-VIII not deflected
ventrad, not forming an anal plate; male genitalia
may be modified, but without a long sword-like
processi: Microvelia Westwood
Tanyvelia gen. n.
(figs. 1-4, 27)
Diagnosis
Size. — Micropterous form, length of males 2.73-
2.91 mm, females 2.81-3.13 mm; general body char-
acteristics and size sexually dimorphic, males smaller,
female abdomen more robust.
Colour. — Ground colour blackish brown, tinged
with orange brown, without silvery pubescence; ante-
rior pronotal lobe yellowish, pruinose, sharply demar-
cated from dark collar and posterior lobe.
Micropterous wing pads brown, set with anterolater-
al light streaks.
Structural characters. — Only micropterous form
known, shape elongate (fig. 1). Eyes globose, exserted
but usually not beyond anterolateral pronotal angles,
separated by about twice the width of an eye, ap-
pressed to anterior pronotal margin, with short ocular
setae. Head declivant anteriorly, recessed into prono-
tum, posterior margin sloping caudo-dorsally, with
usual three pairs of facial trichobothria; gular region
moderately long, plainly visible, rostral cavity closed
posteriorly. Rostrum reaching to middle of mesoster-
num, segment I short, enclosed in rostral cavity, seg-
ments I and IV subequal in length and about three
times longer than II, segment II about 8 times as
long as II. Antennae slender, very long, about 3/4 of
body length.
Pronotum slightly raised medially, with weak me-
dian longitudinal carina; collar weakly formed, set off
by a ragged row of small dark foveae, terminating
under eyes laterally; anterior and posterior lobes set
off by a transverse row of small foveae, evanescent
medially; anterior lobe with scattered stiff erect dark
setae; posterior lobe with numerous small foveae, hu-
meri not prominent, broadly rounded posteriorly,
not modified, covering metanotum. Thoracic venter
not diagnostic, with weakly formed tubercles on ei-
ther side of mesosternal midline on posterior margin
opposing an unmodified metasternum. Metasternal
scent gland opening (omphalium) small but visible,
58
marked by a small tubercle; scent channels promi-
nent, curving slightly anterad to base of metacetabu-
lae.
Abdomen without silvery setae or other adorn-
ment, except weak longitudinal pruinose line on mid-
line of tergites IV-VII; with short paired longitudinal
carinae on basal half of tergite II. Abdominal sternites
set off from laterosternites by hair-free glabrous oval
lacunae. Micropterous wing pads elongate.
Legs slender, long, hind legs very long; anterior fe-
mur set beneath with short light setae, unmodified in
males; anterior tibia of male with a comb of minute
black setae occupying 3/4 of tibial length; middle fe-
mora set ventrally with 4-5 very long slender setae; all
tarsi long (fig. 3), claws moderately long; both up-
and down curving arolia large, evident.
Male genital segments moderately large, protrud-
ing, modified; proctiger unmodified (fig. 4); para-
meres small, slender symmetrical, acuminate (fig. 2).
Female tergite VIII on same plane as VII, truncate
posteriorly; first gonocoxae small, barely exposed,
plate-like; tergite IX of both sexes triangular, protrud-
ing posteriorly.
Type-species: Tanyvelia missim sp. n.
Remarks
Comparative notes. — Separable from other genera
of Papuan Microveliinae by the characters given in
the key. Similar in general appearance and ecological
habits to Baptista Distant, but distinguished by the
presence of a long grasping comb that runs nearly the
entire length of the male fore tibia (fig. 3), and the ab-
sence of modifications to abdominal sterna VI and
VII. Among the Papuan genera, Tanyvelia appears
most closely allied to Aegilipsicola, but the pronotal
collar is not as pronounced, the legs, although long
and slender, are slightly stouter, and the fore tibial
grasping comb is shorter. The proctiger of Tanyvelia
is broadly rounded distally rather than coming to a
rounded point as in Aegilipsicola (compare figs. 4 and
7), and the paramere is slender and elongate rather
than short and stout (compare figs. 2 and 8).
Etymology. — The generic name 7anyvelia is de-
rived from tany- (Gr.), long, referring to the slender
body shape, and Velia, the nominate genus of the
family. Gender feminine.
Distribution. — New Guinea (fig. 27).
Tanyvelia missim sp. n.
(figs. 1-4, 27)
Type material: Holotype, micropterous male:
Papua New Guinea, Morobe Prov., upper Poverty
Creek on Mt. Missim, nr. Wau, 1600 m., 18 Sept.
POLHEMUS & POLHEMUS: Microveliinae from New Guinea
Praga 2 60000
Du u
vv
Figs. 1-4. Tanyvelia missim, gen. n., sp. n. .— 1. Male, dorsal habitus; 2. Male foreleg; 3. Male paramere; 4. Male proctiger.
Figs. 5-8. Aegilipsicola rapida, gen. n., sp. n. — 5. Male, dorsal habitus; 6. Male foreleg; 7. Male proctiger; 8. Male paramere
52)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
1983, CL 1830, D. A. and J. T. Polhemus (BPBM). -
Paratypes: 23 micropterous males, 12 micropterous
females, same data as holotype (JTPC).
Diagnosis
Size. — Micropterous male, length 2.73-2.91 mm
(x = 2.81, n = 10); width 0.79-0.90 mm (x = 0.86, n
= 10). Micropterous female, length 2.81-3.13 mm (x
= 2.96, n = 10); width 0.83-0.94 mm, (x = 0.88, n =
10).
Colour. — Micropterous male: ground colour
black, venter and connexiva tinged with brown. Head
black, brown ventrally; rostrum luteous on basal three
segments, piceous distally. Pronotum with anterior
lobe entirely yellowish including propleura, except
narrowly embrowned medially; disc and collar black.
Abdomen black, tergites mostly pruinose, lighter ven-
trally. Legs and antennae luteous to yellow brown,
distally darker.
Structural characters. — Micropterous male: head
of moderate length, declivant anteriorly; length 0.47;
width of eye/interocular space, 0.16/0.30. Pronotum
length:width, 0.74 : 0.81. Abdominal tergites not
shining; tergites, II-VI subequal in length (0.16-
0.19), VII longer (0.30). Abdominal venter set with
short appressed setae; ventrite VII with a short V-
shaped depression on caudal half. Legs, antennae
thickly clothed with short to moderate length setae,
with scattered longer setae. Legs unarmed.
Antennal formula, segments I : II : HI: IV; 0.51:
0.40 : 0.70 : 0.70.
Proportions of legs as follows: Femur, tibia, tarsal
1, tarsal 2 of fore leg, 0.79 : 0.77 : 0.30 : 0.0; of mid-
dle leg, 1.07 : 1.05 : 0.14 : 0.28; of hind leg, 1.16:
1.47 : 0.23 : 0.30.
First genital segment with an antero-ventrally di-
rected tuft of stiff dark setae. Paramere small, long,
triangular (fig. 2).
Micropterous female: Similar to male in most re-
spects, but somewhat larger; connexiva vertical or re-
flexed over dorsum except in gravid specimens.
Remarks
Comparative notes. — Recognized among the Pa-
puan Microveliinae by its elongate form with long
legs and antennae (fig. 1), relatively unmodified male
foreleg with a long tibial grasping comb (fig. 3), acu-
minate paramere (fig. 2), and elongate male proctiger
(fig. 4).
Biological notes. — Upper Poverty Creek at the Mt.
Missim type locality was a small, rocky headwater
stream dropping steeply through a boulder-strewn
bed heavily shaded by primary rain forest.
Etymology. — The name ‘missim is a noun in appo-
60
sition and refers to the Mt. Missim type locality.
Distribution. — Eastern New Guinea (fig. 27).
Aegilipsicola gen. n.
(figs. 5-8, 28)
Diagnosis
Size. — Macropterous form, length of males 2.70-
2.84 mm, females 3.16-3.31 mm, general body char-
acteristics not sexually dimorphic, but males slightly
smaller.
Colour. — Macropterous form: ground colour
blackish brown, tinged with yellowish brown; dorsal-
ly dark brown on head and ventrally, wings black,
brown to black beneath; anterior pronotal lobe nar-
rowly yellowish on either side of midline, laterally
dark but covered with silvery pubescence except me-
dially, demarcated from dark collar and posterior
lobe. Hemelytra black, bearing six elongate bluish-
grey pruinose streaks, with two confined within the
basal cells, and four others on distal third (fig. 5)
Structural characters. — Only macropterous form
known. Form elongate, slender (fig. 5); eyes globose,
exserted beyond anterolateral pronotal angles, separ-
ated by about twice the width of an eye, removed
from anterior pronotal margin, with short ocular se-
tae. Head moderately declivant anteriorly, recessed
into pronotum, posterior margin sloping somewhat
caudo-dorsally, with usual three pairs of facial tricho-
bothria; gular region long, plainly visible, rostral cav-
ity closed posteriorly by a strong carinate ridge.
Rostrum reaching to middle of mesosternum, seg-
ment I short, enclosed in rostral cavity, I and IV sub-
equal in length and about three times longer than II,
segment III about 8 times as long as II. Antennae very
slender, very long, about 2/3 of body length.
Pronotum slightly raised medially, with weak median
longitudinal carina marked by a line of golden pubes-
cence; collar prominent, distinctly set off by a row of
deep foveae, terminating laterally; near ventral eye
margins; anterior and posterior lobes set off by a
transverse row of foveae, evanescent medially; entire
pronotum sparsely set with fine decumbent golden
pubescence; posterior lobe with numerous obscure
shallow foveae, humeri moderately prominent, al-
most triangular, narrowly rounded posteriorly.
Thoracic venter not diagnostic, with weakly formed
tubercles on either side of mesosternal midline along
posterior margin opposing an unmodified metaster-
num. Metasternal scent gland opening (omphalium)
not visible; scent channels prominent, curving ante-
rad to base of metacetabulae.
Abdomen without silvery setae or other adorn-
ment; prominent paired longitudinal carinae present
on basal half of tergite II in macropters (visible only
after removal of wings). Abdominal sternites set off
from laterosternites by hair-free glabrous oval lacu-
nae. Female connexiva expanded, plate like along ter-
gite VII. Macropters with five closed cells in hemely-
tra, venation of fore and hind wings similar to figs.
312-313 in Andersen (1982) except distal cell closed;
fore wings set with fine semi-recumbent stiff setae on
basal half; micropterous form unknown.
Legs slender, long, hind legs very long; anterior fe-
mur set beneath with short light setae, with ventral
tubercle at distal 2/3 in males; anterior tibia of male
with a fine line of minute black setae occupying 4/5
of tibial length; femora ventrally without long setae;
all tarsi long (fig. 6), claws very long; both up- and
down curving arolia long, slender.
Male genital segments moderately large, protrud-
ing, modified; proctiger unmodified (fig. 7); para-
meres small, slender, symmetrical (fig. 8). Female ter-
gite VIII on same plane as VII, protruding posteriorly
over tergite IX; first gonocoxae small, barely exposed,
plate-like; tergite IX of both sexes button-like, pro-
truding posteriorly.
Type-species: Aegilipsicola rapida sp. n.
Remarks
Etymology. — The generic name Aegilipsicola is de-
rived from Aegilips (Gr.), sheer, steep, referring to the
habitat, and -cola, (L.), inhabitant. Gender feminine.
Comparative notes. — In general facies Aegilipsicola
most closely resembles the Neotropical genus Aegili-
psivelia, however this similarity is apparently due to
convergence. Although the two genera share many
similar structures adapted to life on steep hygropetric
habitats, such as long legs and extremely long claws,
Aegilipsivelia lacks the carinate rostral cavity on the
gula, the bright pruinose hemelytral markings, and
the carinate female connexiva seen in Aegilipsicola. In
addition, the collar in Aegzlipsivelia is set off by a sul-
cus rather than a row of deep foveae, and the comb-
like process on the male fore tibia is relatively short
instead of almost as long as the tibia.
Among Papuan genera of Microveliinae Aegilipsi-
cola is apparently most closely allied to Tanyvelia gen.
n. (see discussion under 7anyvelia)
Distribution. — Eastern New Guinea (fig. 28).
Aegilipsicola rapida sp. n.
(figs. 5-8, 28)
Type material: Holotype, macropterous male:
Papua New Guinea, Morobe Prov., seeps along Mt.
Kaindi road nr. Kunai Creek, above Wau, 17 Sept.
1983, CL 1823, D. A. and J. T. Polhemus (BPBM). —
Paratypes: 39 macropterous males, 37 macropterous
females, 62 immatures, same data as holotype (JTPC);
POLHEMUS & POLHEMUS: Microveliinae from New Guinea
4 macropterous males, 4 macropterous females, 2 im-
matures, Morobe Prov., Namie Creek, Mt. Kaindi,
17 Seas OS SEITE IE SHD 2 Polhemus
(TPC); 1 macropterous female, Morobe Prov., Kunai
River, 1500 m, 10 Oct. 1966, Illies (ZBSM).
Diagnosis
Size. — Macropterous male, length 2.70-2.84 mm
(x = 2.74, n = 10); width 0.86-0.97 mm (x= 0.94, n
= 10). Macropterous female, length 3.06-3.31 mm (x
= 3.16, n = 10); width 1.01-1.12 mm, (x = 1.04, n =
10).
Colour. — Macropterous male: ground colour
black, tinged with yellowish brown ventrally. Head
black; two longitudinal regions between eyes and me-
dial furrow, posterior margin, brown. Pronotum with
anterior lobe narrowly, transversely orange brown on
either side of midline, extending to inner eye margins;
posterior lobe uniformly black. Hemelytra blackish
brown, long, extending beyond tip of abdomen. Ab-
domen brownish yellow ventrally, with a broad
brown longitudinal stripe along sternal-laterosternal
boundary. Legs yellowish, darker distally; antennae
brown, segment I yellowish basally.
Structural characters. — Macropterous male: head
of moderate length, declivant anteriorly; length 0.51;
width of eye/interocular space, 0.14/0.30. Pronotum
long, humeri evident but not raised; length : width,
0.84 : 0.91.
Abdominal venter set with very short appressed
fine setae; venter VII depressed on either side of mid-
line; first genital segment excavated ventrally, with
three small brown sclerotized denticles distally. Legs,
antennae thickly clothed with short inconspicuous se-
tae, without longer setae. Legs unarmed, except fore
femur set near distal 2/3 with patch of short stiff setae
(fig. 6).
Antennal formula I: IL: III : IV; 0.40 : 0.28 : 0.74
: 0.60. |
Proportions of legs as follows: Femur, tibia, tarsal
1, tarsal 2 of fore leg, 0.74 : 0.60 : 0.28 : 0.0; of mid-
dle leg, 1.00 : 0.98 : 0.12 : 0.33; of hind leg, 1.44 :
1.74 : 0.09 : 0.40.
Paramere small, short, ovate (fig. 8).
Macropterous female: Similar to male in most re-
spects, but somewhat larger. Connexiva vertical,
strongly raised and plate-like along tergite VII, pro-
duced posteriorly along tergite VIII.
Remarks
Comparative notes. — Easily recognized among the
Papuan Microveliinae by its elongate form (fig. 5),
broadly pointed proctiger (fig. 7), short and broad pa-
ramere (fig. 8), elongate fore tibial grasping comb
61
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 9-16. Tarsovelia gen. n.—9. Tarsovelia alta sp. n., micropterous female, dorsal habitus; 10. Tarsovelia dani sp. n., male proc-
tiger; 11-13. Male forelegs of Tarsovelia species, 11. Tarsovelia arfak sp. n.; 12. Tarsovelia dani sp. n.; 13. Tarsovelia alta sp. n.;
14-16. Male parameres of Tarsovelia species. 14. Tarsovelia dani sp. n.; 15. Tarsovelia alta sp. n.; 16. Tarsovelia arfak sp. n.
62
(fig. 6), and small tuft of black setae ventrally on the
male fore femur (fig. 6).
Biological notes. — The type series was taken from
wet seeping vertical rock faces formed by road cuts
along the Mt. Kaindi road. The insects were abun-
dant, and ran across the wet rocks and moss with
great speed and agility.
Etymology. — The name ‘rapida refers to the speed
with which these insects can move across vertical rock
faces.
Distribution. — Eastern New Guinea (fig. 28).
Tarsovelia gen. n.
(figs. 9-16, 29)
Diagnosis
Size. — Length of males 2.41-2.95 mm, females
2.77-3.45 mm, general body characteristics and size
sexually dimorphic, males smaller, female abdomen
more robust.
Colour. — Ground colour brown to _ blackish
brown, often blackish above, without silvery pubes-
cence; winged forms dorsally black, wings black,
brown to black beneath. Anterior pronotal lobe and
collar usually entirely yellowish to orange brown, on-
ly in 7: arfak dark laterally.
Structural characters. — Form robust (fig. 9); eyes
globose, exserted but usually not beyond anterolater-
al pronotal angles, separated by at least twice the
width of an eye, appressed to anterior pronotal mar-
gin, with long ocular setae. Head declivant anteriorly,
slightly recessed into pronotum, with usual three
pairs of facial trichobothria; gular region moderately
long, plainly visible, rostral cavity closed posteriorly.
Rostrum reaching almost to metasternum, segment I
short, enclosed in rostral cavity, segments I and IV
subequal in length and about three times longer than
II, segment III about 8 times as long as II. Antennae
slender, long, about 1/2 of body length.
Pronotum of micropterous form slightly raised me-
dially; collar weakly formed, set off by a ragged row of
small dark foveae, terminating under eyes laterally;
anterior and posterior lobes set off by a transverse row
of small foveae in a weak transverse sulcus, evanescent
medially; anterior lobe set with stiff erect setae; poste-
rior lobe with numerous shallow depressions, humeri
not prominent, broadly rounded posteriorly, not
modified, covering metanotum. Pronotum of alate
form longer, rounded posterior margin extending far-
ther caudad, almost triangular, humeri more promi-
nent. Thoracic venter not diagnostic, with weakly
formed tubercles on either side of mesosternal mid-
line on posterior margin opposing an unmodified
metasternum. Metasternal scent gland opening (om-
POLHEMUS & POLHEMUS: Microveliinae from New Guinea
phalium) not evident; scent channels prominent, an-
gled slightly anterad to base of metacetabulae.
Abdomen without silvery setae or other adorn-
ment, except weak longitudinal pruinose line on mid-
line of tergites IV-VII; prominent paired longitudinal
carinae on tergite II of macropters (visible only after
removal of wings), absent in micropterous form.
Abdominal sternites set off from laterosternites by
hair-free round or oval lacunae. Micropters with el-
ongate white wing pads, often infuscated distally;
macropters with four closed cells in dark hemelytra
with white region basally; venation of fore and hind
wings similar to figs. 312-313 in Andersen (1982); fe-
male fore wing Sc often terminating in a triangular
thickened region, resulting in a slight to pronunced
sinuosity of the wing margin; fore wings set with long
dark stiff erect setae on basal half, much more pro-
nounced in females.
Legs slender; anterior femur set beneath with short
light setae, sometimes slightly modified in males (figs.
11-13); anterior tibia with a distal comb of many stiff
setae; middle and anterior femora set ventrally with
numerous long slender setae; middle tibia set with an
evenly spaced row of about 10 erect setae beginning
at basal third, very long basally, decreasing in length
distally; middle tarsi extremely long (fig. 9); claws
short; both up- and down curving arolia evident, but
very slender.
Male genital segments small, not protruding, un-
modified; proctiger unmodified; parameres small,
slender symmetrical (figs. 14-16). Female tergite VIII
on same plane as VII, truncate posteriorly; first gono-
coxae small, barely exposed, plate-like; tergite IX
small, button-like, protruding posteriorly.
Type-species: Tarsovelia alta sp. n.
Remarks
Comparative notes. — Zarsovelia is similar in gener-
al facies to Microvelia, but is easily separated from this
genus and all other Microveliinae by the elongate
middle tarsi (fig. 9), which are adapted for rowing rat-
her than running. In addition, the light colored mar-
kings on the hemelytra of Tarsovelia are restricted to
the basal angles, plus occasionally a weak medial fas-
cia, whereas in the type-species of Microvelia (M. pul-
chella Westwood) the hemelytra also bear ovate light
spots in every closed cell, plus a large bright spot in
the open distal cell.
Biological notes. — Tarsovelia species are found
along the margins of flowing pools on rocky moun-
tain streams. They are swift and agile skaters if dis-
turbed, and will fly readily if captured in a net.
Etymology. — The generic name Tarsovelia is de-
rived from tarsos (Gr.), referring to the extremely long
63
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
slender middle tarsi, and Velia, the nominate genus of
the family. Gender feminine.
Distribution. — New Guinea (fig. 29).
Key to the species of Tarsovelia
Males
1. Fore femur with patch of short, stiff hairs on
ventralisurface((fie si) eee ee 22
— Fore femur lacking patch of short, stiff hairs on
ventralisurfaeei(ti DS): T. alta sp. n.
2. Fore femur with patch of short, stiff hairs near
middle of ventral margin (fig. 11) … nn
TIBIA arte Bal BERE. Leda T. arfak sp. n.
— Fore femur with patch of short, stiff hairs near
base of ventral margin (fig. 12) ..... T. dani sp. n.
Tarsovelia alta sp. n.
(figs. 9, 13, 15, 29)
Type material: Holotype, micropterous male:
Papua New Guinea, Morobe Prov., bog nr. Mt.
Kaindi summit, above Wau, 17 Oct. 1983, CL 1825,
D. A. and J. T. Polhemus (BPBM). — Paratypes: 13 mi-
cropterous males, 14 micropterous females, 16 imma-
tures, same data as holotype (ZMUC, JTPC, BPBM).
Additional material examined: 2 macropterous fe-
males, Papua New Guinea, Morobe Prov., Eddie
Creek, 2050 m, 11 Oct. 1966, Illies leg. (ZBSM, TPC);
1 macropterous female, Morobe Prov., Poverty
Creek, Mt. Missim, 18 Sept. 1983, CL 1830, 1600
m, J. T. & D. A. Polhemus (JTPC); 9 macropterous
males, 6 macropterous females, 2 immatures, West-
ern Highlands Prov., rocky stream 17 km N. of Mt.
Hagen on Baiyer River road, 6 and 8 Sept. 1983, CL
1780, D. A. and J. T. Polhemus (JTPC).
Diagnosis
Size. — Micropterous male, length 2.41-2.48 mm
(x = 2.44, n = 10); width 0.83-0.89 mm (x = 0.85, n
= 10). Micropterous female, length 2.81-2.88 mm (x
= 2.84, n = 4); width 0.94-1.01 mm, (x = 0.99, n = 4).
Macropterous male, length 2.77-2.95 mm (x =
2.87, n = 9); width 1.10-1.15 mm (x = 1.14, n = 9).
Macropterous female, length 3.31-3.45 mm (x =
3.36, n = 6); width 1.19-1.33 mm, (x = 1.28, n = 6).
Colour. — Micropterous male: ground colour
orange brown, venter slightly lighter. Head dark
orange brown, lighter beneath; rostrum luteous medi-
ally, piceous distally. Pronotum with anterior trans-
verse band orange brown, extending onto propleura;
disc dark brown, mottled with lighter brown.
Abdomen orange brown, tergal margins darker, light-
er ventrally. Distal segements of legs, antennae
brown, basal segments lighter.
Structural characters. — Micropterous male: head
64
of moderate length, declivant anteriorly; length 0.37;
width of eye/interocular space, 0.14/0.30. Pronotum
long, covering metanotum, humeri evident but not
raised; length:width, 0.84 : 0.86. Abdominal tergites
not shining; lengths of tergites, II : 0.14; III : 0.16;
IV-VI : 0.19; VII : 0.26.
Abdominal venter not modified, set with short ap-
pressed setae. Legs, antennae thickly clothed with
short to moderate length setae, with scattered longer
setae. Legs unarmed.
Antennal formula I : II: III: IV; 0.29 : 0.33 : 0.37
: 0.42.
Proportions of legs as follows: Femur, tibia, tarsal
1, tarsal 2 of fore leg, 0.65 : 0.63 : 0.28 : 0.0; of mid-
dle leg, 0.98 : 0.93 : 0.42 : 0.35; of hind leg, 0.86 :
1.00 : 0.19 : 0.29.
Paramere small, long, slender (fig. 15).
Micropterous female: Similar to male in most re-
spects, but somewhat larger; connexiva reflexed over
dorsum except in gravid specimens.
Macropterous male: Similar to micropterous form
in most respects. Pronotum with anterior lobe entire-
ly orange brown, extending onto propleura; posterior
lobe uniformly dark. Hemelytra blackish brown, ba-
sally entirely white except slight infuscation of Sc,
white region extending beyond tip of pronotum.
Macropterous female: Very similar to macropter-
ous male, except lateral hemelytral margin slightly
sinuate at distal thickening of Sc. Posterior lobe of
pronotum with two (1+1) tufts of erect stiff black se-
tae on either side of midline at about middle.
Connexival margin vertical.
Remarks
Comparative notes. — 7. alta sp. n. may be separa-
ted from 7. dani sp. n. and 7: arfak sp. n. by the un-
modified male fore femur (fig. 13), brownish colora-
tion in wingless forms, male paramere shape (fig.15),
and length of the second segment of the hind tarsi,
which is about 1.5 times as long as the first (versus
only slightly longer in the other two species). In 7:
arfak sp. n. the anterior femora are dark distally, and
the middle and posterior femora plus all the tibiae
and tarsi are dark brown to black, while in 7. alta sp.
n. and 7. dani sp. n. all femora are largely light
brown, and the tibiae and tarsi are dark brown but
not black. In the macropters of 7. alta sp. n. and 7:
arfak sp. n. the entire basal angle of the hemelytra is
white (including the veins, although these are occa-
sionally slightly darkened), whereas in 7: dani sp. n.
the veins are dark and the light coloring is often a sor-
did white.
We originally considered the micropterous and
macropterous specimens at hand from Papua New
Guinea to represent different taxa. However with two
additional new species now available from Irian Jaya
that show marked differences from the Papua New
Guinea material, we have now concluded that the lat-
ter represent but a single, albeit somewhat variable,
species. We have thus included all of the specimens
from eastern New Guinea under 7. alta sp. n., but the
macropters are not named as paratypes. Macropte-
rous specimens from Eddie Creek and Poverty Creek
(not measured) have very long hemelytra, consider-
ably exceeding the tip of the abdomen, and they are
lighter in color than specimens from Mt. Hagen.
Measurements of leg and antennal segments are not
helpful, as they appear to vary randomly between
populations, thus we defer the final decision about
the specific distinctness of these populations from
Papua New Guinea until more material, hopefully in-
cluding both micropters and macropters in the same
sample, is available.
Biological notes. — Near Mt. Hagen 7. alta was
found in a plunge pool below a small cascade, with
individuals hiding in the crevices of a sheer rock wall.
When disturbed the insects would skate rapidly over
the water near the wall until disappearing into their
hiding places once more. By contrast, the Mt. Kaindi
specimens were taken from beneath a large, partly
submerged log in a bog pool, where violent flushing
was required to dislodge them. So far this species has
been found only at altitudes above 1200 meters.
Etymology. — The name ‘a/ta’ refers to the high el-
evation at which this species was taken.
Distribution. — Eastern New Guinea (fig. 29).
Tarsovelia arfak sp. n.
(figs. 11, 16, 29)
Type material: Holotype, macropterous male:
Indonesia, Irian Jaya Prov., Saumarin River, nr.
Warkomi, Arfak Mountains, 42 km S. of
Manokwari, 90 m, water temp eG. 18 Oct 1991,
CL 2647, D. A. and J. T. Polhemus (USNM). —
Paratypes: 67 macropterous males, 48 macropterous
females, 6 immatures, same data as holotype (JTPC).
Diagnosis
Size. — Macropterous male, length 2.45-2.77 mm (x
= 2.62, n = 10); width 1.01-1.12 mm (x = 1.06, n =
10). Macropterous female, length 2.81-3.09 mm (x =
2.98, n = 10); width 1.15-1.19 mm, (x = 1.17, n= 10).
Colour. — Macropterous male: ground colour
black, tinged with brown. Head black. Pronotum
with anterior lobe and collar medially orange brown,
extending to inner eye margins; posterior lobe uni-
formly black. Hemelytra blackish brown, long, ex-
tending beyond tip of abdomen, basally with broad
elongate rectangular white region along outer margin,
POLHEMUS & POLHEMUS: Microveliinae from New Guinea
extending beyond tip of pronotum. Abdomen black,
ventrally tinged with brown along connexival mar-
gins and caudally. Legs, antennae brown, fore femur
yellowish except distally, mid and hind femora yel-
lowish ventrally.
Structural characteristics. — Macropterous male:
head of moderate length, declivant anteriorly; length
0.40; width of eye/interocular space, 0.12/0.33.
Pronotum long, humeri evident but not raised;
length:width, 0.86 : 1.00.
Abdominal venter not modified, set with short ap-
pressed setae. Legs, antennae thickly clothed with
short to moderate length setae, with scattered longer
setae; all tibia, middle and hind tarsi set with rows of
long setae dorsally, in addition to long ventral setae.
Legs unarmed, except fore femur ventrally slightly ex-
cavate basally, set near middle with patch of short stiff
setae (fig. 11).
Antennal formula I: II: III: IV; 0.28 : 0.33 : 0.35
: 0.40.
Proportions of legs as follows: Femur, tibia, tarsal
1, tarsal 2 of fore leg, 0.65 : 0.51 : 0.28 : 0.0; of mid-
dle leg, 0.88 : 0.81 : 0.49 : 0.35; of hind leg, 0.81 :
0.95 : 0.23 : 0.28.
Paramere small, long, slender (fig. 16).
Macropterous female: Similar to male in most re-
spects, but somewhat larger; hemelytra with lateral
margin sinuate, thickened and set with long dense
black setae at distal Sc. Connexiva vertical along ter-
gites II-IV, slightly reflexed over tergites V-VII.
Remarks
Comparative notes. — Recognized by the patch of
short stiff black setae on the ventral margin of the ma-
le fore femur (fig. 11), and the shape of the male pa-
ramere (fig. 16). For additional comparative notes see
key, and discussion under 7. alta sp. n.
Biological notes. — The Saumarin River at the type
locality was a moderate sized stream dropping steeply
to the sea off the flanks of the Arfak Mountains
through a bed of boulders, cobbles and sand, heavily
shaded by primary rain forest. The stream profile was
punctuated by numerous small waterfalls, and 7: ar-
fak was most abundant on the plunge pools below
these falls.
Etymology. — The name ‘arfak’ is a noun in appo-
sition and refers to the Arfak Mountain type area.
Distribution. — Western New Guinea (Vogelkop
Peninsula) (fig. 29).
Tarsovelia dani sp. n.
(figs. 10, 12, 14, 29)
Type material: Holotype, macropterous male:
Indonesia, Irian Jaya Prov., swift rocky stream in
65
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
66
upper Pass Valley, 52 km NE of Wamena, 2015 m.,
water temp. 14°C, 23 Sept. 1991, CL 2614, D. A.
and J. T. Polhemus (USNM). - Paratypes: 61 macrop-
terous males, 26 macropterous females, 28 imma-
tures, same data as holotype (JTPC).
Diagnosis
Size. — Macropterous male, length 2.55-2.95 mm
(x = 2.78, n = 10); width 1.01-1.15 mm (x = 1.09, n
= 10). Macropterous female, length 2.99-3.45 mm (x
= 3.12, n = 10); width 1.15-1.37 mm, (x = 1.22, n =
10).
Colour. — Macropterous male: Ground colour
orange brown, marked with black. Head blackish
brown dorsally, ventrally lighter. Pronotum with ante-
rior lobe and collar orange brown, extending onto
propleura, collar marked only laterally, demarcated by
a row of dark foveae; posterior lobe uniformly black.
Hemelytra blackish brown, basally with narrow elon-
gate white stripe along costal margin, extending be-
yond tip of pronotum, plus a shorter sordid white re-
gion basally along pronotum; with elongate grayish
brown streak in middle, barely visible in some speci-
mens; length variable, shortest reaching barely onto
tergite VII, longest almost covering genital segments.
Abdomen orange brown. Legs, antennae brown, fore
femur yellowish except distally, mid and hind femora,
tibia yellowish ventrally.
Structural characteristics. — Macropterous male:
head of moderate length, declivant anteriorly; length
0.38; width of eye/interocular space, 0.14/0.30.
Pronotum long, humeri evident but not raised;
length:width, 1.00 : 1.12.
Abdominal venter not modified, set with short ap-
pressed setae. Legs, antennae thickly clothed with
short to moderate length setae, with scattered longer
setae. Legs unarmed, except fore femur flattened ven-
trally, curved, basally with a tuft of stiff setae (fig. 12).
Antennal formula I : II : IMI : IV; 0.30 : 0.35 : 0.42
: 0.42.
Proportions of legs as follows: Femur, tibia, tarsal
1, tarsal 2 of fore leg, 0.74 : 0.72 : 0.30 : 0.0; of mid-
dle leg, 1.12 : 1.02 : 0.49 : 0.35; of hind leg, 0.98 :
1.07 : 0.26 : 0.33.
Paramere small, long, slender (fig. 14).
Macropterous female: Similar to male in most re-
spects, but somewhat larger. Hemelytra length vari-
able, shorter than in male, shortest not reaching ter-
gite VII, longest reaching caudal margin of tergite
POLHEMUS & POLHEMUS: Microveliinae from New Guinea
VII; with lateral margin weakly sinuate, narrowly
thickened and set with stiff setae at distal Sc.
Connexiva vertical along tergites II-III, IV in transi-
tion, strongly reflexed over tergites V-VII.
Remarks
Comparative notes. — Recognized by the small tuft
of black setae basally on the ventral surface of the we-
akly bowed male fore femur (fig. 12), and the shape
of the male paramere (fig. 14). For additional compa-
rative notes see key, and discussion under 7: alta sp.
n.
Biological notes. — The type series of 7. dani was
taken from pools along a very swift, cold mountain
stream in a limestone bed bordered by Nothofagus and
Rhododendron forest. The insects were most common
along the pool margins, skating rapidly across the wa-
ter if disturbed and flying readily if taken in a net.
Etymology. — The name ‘dan? is a noun in apposi-
tion and refers to the native Dani people who inhabit
the type area.
Distribution. — Central New Guinea (fig. 29).
Neusterinsifer gen. n.
(figs. 17-26, 30)
Diagnosis
Size. — Form stout (fig. 17), length of males 1.87-
2.48 mm, females 1.91-2.77 mm, general body char-
acteristics and size sexually dimorphic, males smaller,
female abdomen more robust.
Colour. — Ground color black, tinged with orange
brown, heavily marked with silvery pubescence; ante-
rior pronotal lobe orange brown on posterior half,
with streak extending from near midline to middle of
eyes, weakly demarcated from collar and posterior
lobe.
Structural characteristics. — Both apterous and
macropterous forms known. Eyes globose, exserted
but usually not beyond anterolateral pronotal angles,
separated by about twice the width of an eye, ap-
pressed to anterior pronotal margin, with moderately
long ocular setae. Head strongly declivant anteriorly,
recessed into pronotum, posterior margin almost ver-
tical, with usual three pairs of facial trichobothria; gu-
lar region short, not visible, rostral cavity open poste-
riorly, with raised carinate bucculae. Rostrum
reaching to middle of mesosternum, segment I short,
enclosed in rostral cavity, I and IV subequal in length
Figs. 17-26. Neusterinsifer gen. n. — 17. Neusterinsifer compactus sp. n., micropterous male, dorsal habitus; 18-22. Processes
arising from male proctigers of Neusterinsifer species, 18. Neusterinsifer compactus sp. n.; 19. Neusterinsifer cyclops sp. n.; 20.
Neusterinsifer sepik sp. n.; 21. Neusterinsifer nabire sp. n.; 22. Neusterinsifer gladius sp. n.; 23. Neusterinsifer compactus sp. n.,
posterior view of female abdomen showing anal plate formed by tergites VII and VIII. 24-26. Male forelegs of Neusterinsifer
species: 24. Neusterinsifer sepik sp. n.; 25. Neusterinsifer gladius sp. n.; 26. Neusterinsifer compactus sp. n.
67
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
and about two times longer than II, segment III
about four times as long as II. Antennae slender, very
long, about 2/3 of body length.
Pronotum almost flat, without median carina; col-
lar very weakly formed, set off by a ragged row of
small dark foveae, terminating under eyes laterally;
anterior and posterior lobes set off by a transverse row
of foveae; anterior lobe thickly set with appressed sil-
very setae except triangular median area; posterior
lobe with numerous small foveae, humeri not formed,
almost straight posteriorly, not modified, covering
mesonotum. Metanotum almost as long as prono-
tum. Thoracic venter not diagnostic, with weakly
formed tubercles on either side of mesosternal mid-
line on posterior margin opposing an unmodified
metasternum. Metasternal scent gland opening (om-
phalium) not visible, but marked by a tiny tubercle;
scent channels prominent, curving slightly anterad to
base of metacetabulae.
Abdomen with silvery setae laterally on tergites II,
VII, VII, covering most of tergites II-VI in females,
less extensive in males; prominent paired longitudinal
carinae on tergite II of macropters (visible only after
removal of wings), absent in micropterous form.
Abdominal sternites set off from laterosternites by
small hair-free glabrous oval lacunae; laterosternite
VII of both sexes set with small round tuft of stiff pos-
terolaterally directed setae, near connexival margin at
basal 2/5 in males, much closer to base in females.
Macropterous forms with four closed cells in dark
hemelytra, occasionally with fifth distal cell closed; al-
ways with white elongate streak basally in outer basal
cell, usually with faint to pronounced light areas in
other cells; venation of fore and hind wings similar to
figs. 312-313 in Andersen (1982).
Legs moderately stout, moderately long; anterior
femur set beneath with short light setae, modified in
males of some species, with a patch of stiff setae or
denticles ventrally on basal half; anterior tibia of male
with a comb of minute black setae occupying 2/5 to
1/2 of tibial length; all femora set ventrally with 7-12
long slender setae; all tarsi moderately long, claws
moderately long; both up- and down curving arolia
long, slender.
Male abdominal ventrites VI, VII modified; genital
segments very large, protruding, highly modified,
proctiger bearing a large sword-like appendage arising
from anterior margin, directed antero-laterally be-
neath abdomen (figs. 18-22); segment IX, proctiger
rotated by about 90°; parameres vestigial or absent.
Female tergite VIII on same plane as VII basally, dis-
tally sharply deflected ventrad closing abdomen pos-
teriorly except for protruding tip of proctiger (fig.
23); first gonocoxae hidden.
Type-species: - Neusterinsifer compacta sp. n.
68
Remarks
Comparative notes. — Very similar in general facies
to Microvelia (fig. 17), but separated by sword-like
projection on the male proctiger (figs. 18-22), the
small tuft of stiff setae on laterotergite VII, and other
characters as given in the generic key. Macropters
differ from Microvelia and most other genera of
Microveliinae in having a well defined long slender
white streak basally in the outer cell (with only ill de-
fined light areas in all other cells), and lacking the dis-
tal white spot that is characteristic of Microvelia and
other closely related genera. The wing markings are
similar to those of Tenagovelia Kirkaldy from Africa,
but species in this latter genus are much larger and
possess much longer legs.
Biological notes. — Species of Neusterinsifer are
most commonly encountered in sheltered areas along
the margins of stream pools, or on the surfaces of wa-
ter filled potholes in bedrock exposures adjacent to
flowing sections of streams.
Etymology. — The generic name Neusterinsifer is
derived from neuster (Gr.), swimmer, and ensifer (L.),
sword bearer. Gender masculine.
Distribution. — New Guinea (fig. 30).
Key to species of Neusterinsifer
1. Male process on proctiger bifurcate, short (fig.
21). Female abdomen strongly narrowed posteri-
Orlysboatishaped per nn rer nabire sp. n.
— Male process on proctiger not bifurcate, longer (-
figs. 18-20, 22). Female abdomen not strongly
narrowed posteriorly, not boat shaped ............ 2.
2. Small species, male 1.87 mm, female 1.91 mm
long. Male process on proctiger relatively short,
broad, narrowly triangular distally (fig. 19 .........
aan neeh cyclops sp. n.
— Larger species, male at least 2.19 mm, female at
least 2.27 mm long. Male process on proctiger
long, relatively narrow, not triangular distally (-
D 3
3. Relatively small species, males 2.19-2.27 mm, fe-
males 2.27-2.34 mm long. Process of male proc-
tiger long, broad, expanded medially and distally
(fig. 22); male fore femur with a row of 5-10
black spinules basally (fig. 25). Female connexival
segment VII forming a small acute angle at level
of caudal margin of tergite VII ...... gladius sp. n.
— Larger species, males 2.27-2.48 mm, females
2.37-2.77 mm long. Process of male proctiger
long, narrow or broad, not expanded medially
and distally (figs. 18, 20); male fore femur with-
out a basal row of 5-10 black spinules, instead
with either a basal row of many short stiff black
setae, or unarmed. Caudal extreme of female con
nexival segment VII either forming an acute or
obtuse angle, but without a small acute angle at
level of caudal margin of tergite VII … … … … 4
4. Process of male proctiger long, slender, sharp dis-
tally (fig. 18); male fore femur strongly bowed,
bearing small black denticles basally (fig. 26).
Caudal extreme of female connexival segment
VII forming an obtuse angle ..... compactus sp. n.
— Process of male proctiger long, broad (fig. 20);
male fore femur not bowed, with a patch of black
spinules basally (fig. 24). Caudal extreme of fe-
male connexival segment VII slightly produced
posteriorly, forming an acute angle ... sepik sp. n.
Neusterinsifer compactus sp. n.
(figs. 17, 18, 23, 26, 30)
Type material: Holotype, apterous male: Papua
New Guinea, Western Highlands Prov., Baiyer River,
53 km N. of Mt. Hagen, 6 and 8 Sept. 1983, D. A.
and J. T. Polhemus (BPBM). - Paratypes: 19 apterous
males, 12 apterous females, same data as holotype
(TPC); 1 apterous male, 3 apterous females, Western
Highlands Prov., Baiyer River Bird Sanctuary, 8 Sept.
1983, CL 1792, J. T. & D. A. Polhemus (JTPC); 1 ap-
terous male, Morobe Prov., Bamboo Creek, nr. Wau,
16 Sept. 1983, CL 1817, J. T. & D. A. Polhemus
(TPC); 4 apterous males, 1 apterous female, Morobe
Prov., trib. to Bumbu River, N. of Lae, 20 Sept.
1983, CL 1836, J. T. & D. A. Polhemus (JTPC).
Diagnosis
Size. — Apterous male, length 2.37-2.48 mm (x =
2.43, n = 10); width 1.33-1.40 mm (x = 1.39, n =
10). Apterous female, length 2.63-2.77 mm (x =
2.70, n = 10); width 1.40-1.55 mm, (x = 1.50, n =
10).
Colour. — Apterous male: Ground colour blackish
brown, venter slightly lighter, entire dorsum, latero-
tergites covered with fine appressed golden pubes-
cence. Head black, orange brown beneath; rostrum
fuscous medially, piceous distally. Pronotum with an-
terior transverse orange brown band barely interrupt-
ed medially; disc blackish brown. Abdomen blackish
brown, lighter ventrally, connexiva margined with
orange brown. Antennae brown; anterior legs yellow-
ish, distally darkened; middle, hind legs dark brown,
basally lighter; coxae, trochanters luteous.
Structural characters. — Apterous male: head of
moderate length, declivant anteriorly, with weak im-
pressed median line; length 0.40; width of eye/inter-
ocular space, 0.15/0.30. Pronotum long, covering
metanotum, thickly set with obscure foveae, humeri
depressed; length:width, 0.40 : 1.00. Metanotum
length:width, 0.33 : 1.00. Abdominal tergites not
POLHEMUS & POLHEMUS: Microveliinae from New Guinea
shining; tergites II-VI subequal in length (0.14-0.16),
VII longer (0.28).
Abdominal venter set with short appressed setae,
and scattered longer setae caudally; venter VI posteri-
orly set with two (1+1) tufts of dense dark erect setae
on either side of midline; venter VII broadly de-
pressed medially, carinate along posterior margin.
Legs, antennae thickly clothed with short to moderate
length setae, with scattered longer setae. Middle and
hind legs unarmed; fore femur strongly bowed, bear-
ing a patch of black denticles basally, fore tibia with
grasping comb extending 3/4 the length of the tibia
(fig. 26).
Antennal formula I: II : III: IV; 0.35 : 0.28 : 0.40
0511
Proportions of legs as follows: Femur, tibia, tarsal
1, tarsal 2 of fore leg, 0.65 : 0.60 : 0.28 : 0.0; of mid-
dle leg, 0.86 : 0.86 : 0.16 : 0.30; of hind leg, 0.93 :
1.07 : 0.17 : 0.30.
Proctiger produced anteriorly into a long slender
sinuate ensiform process (fig. 18). Parameres vestigial
or absent.
Micropterous female: Similar to male in most re-
spects, but somewhat larger; connexiva vertical, in-
curved and slightly refexed at caudal extreme, which
forms an obtuse angle.
Remarks
Comparative notes. — A relatively large species for
the genus, recognized by the short, blunt process on
the male proctiger (fig. 18), and the strongly bowed
male foreleg with a patch of black denticles basally on
the ventral surface (fig. 26). Similar in size and gener-
al facies to N. sepik sp. n., but this latter species pos-
sesses an unbowed foreleg with a large patch of black
denticles (fig. 24), and a much larger and more poin-
ted process on the male proctiger (fig. 20).
Biological notes. — The Baiyer River at the type lo-
cality was a swift, deep river flowing in a rocky bed
with wet mossy banks, and partially shaded by dis-
turbed upland rain forest. N. compactus was taken
here skating on the surface of water filled potholes in
an exposure of igneous rock along the river margin.
Etymology. — The name compactus (L.) refers to the
shape of this species.
Distribution. — Eastern New Guinea (fig. 30).
Neusterinsifer cyclops sp. n.
(figs. 19, 30)
Type material: Holotype, apterous male: INDONE-
SIA, Irian Jaya Prov., small rocky stream and spring
above Sentani, Cyclops Mountains, 325 m, 25 Sept.
1991, CL 2618, J. T. & D. A. Polhemus (JTPC, to be
placed in USNM when more material is available).
69
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Diagnosis
Size. — Apterous male, length, x = 1.87, n = 1;
width, x = 0.79, n = 1. Apterous female, length, x =
1.91, n = 1; width, x = 0.90, n= 1.
Colour. — Apterous male: Ground colour deep
brown, marked with brown, venter slightly lighter,
entire dorsum, laterotergites covered with fine ap-
pressed golden pubescence; entire dorsum, pleura,
sides of abdomen covered with moderate length erect
setae. Head brown, orange brown beneath; rostrum
fuscous medially, piceous distally. Pronotum with an-
terior transverse orange brown band embrowned me-
dially; disc blackish brown. Abdomen blackish
brown, lighter ventrally, connexiva margined with
orange brown. Antennae brown, base of segment I
lighter; legs yellowish, distally darkened; femora dark-
er posteriorly; coxae, trochanters luteous.
Structural characters. — Apterous male: head of
moderate length, declivant anteriorly, with weak im-
pressed median line; length 0.33; width of eye/inter-
ocular space, 0.12/0.30. Pronotum long, covering
metanotum, thickly set with obscure foveae, humeri
depressed; length:width, 0.28 : 0.79. Metanotum
length:width, 0.26 : 0.79. Abdominal tergites not
shining; tergites II-VI subequal in length (0.12-0.14),
VII longer (0.26).
Abdominal venter set with short appressed setae;
venter VI posteriorly set with two (1+1) tufts of dense
dark erect setae on either side of midline, between
them a V-shaped depression open posteriorly; venter
VII broadly depressed medially, sharply excavated on
posterior half, carinate along anterior margin of exca-
vation. Legs, antennae thickly clothed with short to
moderate length setae, with scattered longer setae.
Middle and hind legs unarmed; foreleg lacking denti-
cles or spinules, tibial comb extending for only 1/2
the length of the tibia.
Antennal formula I: II: III: IV; 0.21 : 0.19 : 0.33
: 0.40.
Proportions of legs as follows: Femur, tibia, tarsal
1, tarsal 2 of fore leg, 0.51 : 0.44 : 0.19 : 0.0; of mid-
dle leg, 0.63 : 0.60 : 0.12 : 0.19; of hind leg, 0.74 :
0.86 : 0.12 : 0.26.
Proctiger produced anteriorly into a broad ensif-
orm process (fig. 19). Parameres vestigial or absent.
Apterous female: Similar to male in most respects,
but somewhat larger; connexiva vertical; caudal ex-
treme gently curving ventrad.
Macropterous forms: unknown.
Remarks
Comparative notes. — Separable from all other
Neusterinsifer species by its small size, and short, poin-
ted process on the male proctiger (fig. 19).
70
Biological notes. — The type of N. cyclops was taken
from a small, clear, spring fed stream at the margin of
the remaining primary rain forest on the slopes of the
Cyclops Mountains above Sentani. The stream
flowed in a bed of metamorphic rocks intermixed
with coarse grey sand, and was partially shaded by
trees felled for garden clearings.
Etymology. — The name cyclops, a noun in apposi-
tion, refers to the mountain range where the species
was found.
Distribution. — Northern New Guinea (Cyclops
Mountains) (fig. 30).
Neusterinsifer gladius sp. n.
(fig. 22, 25, 30)
Type material: Holotype, apterous male: INDONE-
SIA, Irian Jaya Prov., Nabire River and swift tributary,
5 km E. of Nabire, 60 m, 14 Oct. 1991, CL 2642, J.
T. & D. A. Polhemus (USNM). - Paratypes: 2 apterous
males, 2 apterous females, same data as holotype
(JTPC).
Diagnosis
Size. — Apterous male, length 2.19-2.27 mm (x =
2.24, n = 3); width 0.90-0.94 mm (x = 0.91, n = 3).
Apterous female, length 2.27-2.34 mm (x= 2.30, n =
2); width 1.01-1.04 mm, (x = 1.03, n = 2).
Colour. — Apterous male: Ground colour blackish
brown, venter slightly lighter, entire dorsum, latero-
tergites covered with fine appressed golden pubes-
cence; entire body except thoracic venter covered
with long erect setae. Head black, orange brown be-
neath; rostrum fuscous medially, piceous distally.
Pronotum with anterior transverse orange brown
band sometimes interrupted medially; disc blackish
brown. Abdomen blackish brown, lighter ventrally,
connexiva margined with orange brown. Antennae
brown; anterior femora luteous, distally darkened;
anterior tibia, tarsi brown, darker distally; middle,
hind legs brown dorsally, ventrally yellowish, femora
basally lighter; coxae, trochanters luteous, tinged with
fuscous.
Structural characters.- Apterous male: head of mo-
derate length, declivant anteriorly, with weak im-
pressed median line; length 0.35; width of eye/inter-
ocular space, 0.15/0.28. Pronotum long, covering
metanotum, thickly set with obscure foveae, humeri
depressed; length:width, 0.37 : 0.86. Metanotum
length:width, 0.28 : 0.86. Abdominal tergites not
shining; tergites II-VI subequal in length (0.12-0.14),
VII longer (0.28).
Abdominal venter set with short appressed setae;
venter VI posteriorly set with two (1+1) widely separ-
ated tufts of dense dark erect setae on either side of
Fig. 27. Distribution of
Tanyvelia missim, gen. n.,
sp. n.
POLHEMUS & PoLHEMUS: Microveliinae from New Guinea
Fig. 28. Distribution of
Aegilipsicola rapida, gen. n.,
sp. n.
192 134 56
n ni —
New Guinea RTE
o eo ce mo co D
A n A à
i 4 AT
midline, with shallow hair free sulcus between, ex-
tending the length of ventrites V-VI; venter VII
broadly and strongly depressed medially. Legs, anten-
nae thickly clothed with short to moderate length se-
tae, with scattered longer setae. Middle and hind legs
unarmed; fore tibia bearing a row of 5-10 black spi-
nules basally; fore femur with grasping comb extend-
ing 1/2 the length of the tibia (fig. 25).
Antennal formula I: II : III : IV; 0.35 : 0.28 : 0.35
: 0.47.
Proportions of legs as follows: Femur, tibia, tarsal
1, tarsal 2 of fore leg, 0.60 : 0.51 : 0.28 : 0.0; of mid-
dle leg, 0.79 : 0.74 : 0.16 : 0.30; of hind leg, 0.84 :
1.02 : 0.16 : 0.30.
Proctiger produced anteriorly into a long broad
sinuate ensiform process, expanded medially and dis-
tally (fig. 22). Parameres vestigial or absent.
Apterous female: Similar to male in most respects,
but somewhat larger; connexiva vertical; caudal ex-
treme of VII forming an acute angle; VIII gently
curving ventrad.
Macropterous forms: Unknown.
Remarks
Comparative notes. — Similar in size to N. nabire
sp. n., but easily separable by the elongate process on
the male proctiger (fig. 22), the patch of stiff, black
spinules basally on the ventral surface of the male fo-
re femur (fig. 25), and the relatively unmodified fe-
male abdomen which is not laterally compressed or
boat-shaped.
Biological notes. — The Nabire River at the type lo-
cality was a swift, clear, moderate sized stream flow-
ing in a rocky bed, and shaded by primary lowland
rain forest. The type series of N. gladius was taken
downstream of a limestone waterfall, where the banks
of the river included numerous boulders and bedrock
exposures. The insects were taken from the surfaces of
water filled potholes and on still pools amid the rocks.
Etymology. — The name gladius (L.), refers to the
ensiform process of the male proctiger.
Distribution. — Western New Guinea (fig. 30).
Neusterinsifer nabire sp. n.
(figs. 21, 30)
Type material: Holotype, apterous male: INDONE-
SIA, Irian Jaya Prov., Nabire River and swift tributary,
5 km E. of Nabire, 60 m, 14 Oct. 1991, CL 2642, J.
T. & D. A. Polhemus (USNM). - Paratypes: 98 apter-
ous males, 13 macropterous males, 72 apterous fe-
A
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
males, 35 macropterous females, 32 immatures, same
data as holotype (JTPC).
Diagnosis
Size. — Apterous male, length 2.01-2.09 mm (x =
2.04, n = 10); width 0.79-0.83 mm (x = 0.81, n =
10). Apterous female, length 2.19-2.41 mm (x =
2.27, n = 10); width 0.79-0.90 mm (x = 0.83, n =
10). Macropterous male, length 2.23-2.34 mm (x =
2.29, n = 6); width 1.04-1.12 mm (x = 1.10, n = 6).
Macropterous female, length 2.41-2.55 mm (x =
2.46, n = 6); width 1.08-1.15 mm, (x = 1.12, n = 6).
Colour. — Apterous male: Ground colour blackish
brown, venter slightly lighter, entire dorsum, latero-
tergites covered with fine appressed inconspicuous
pubescence; dorsum, body laterally covered with long
erect setae. Head black, orange brown beneath; ros-
trum fuscous medially, piceous distally. Pronotum
with anterior transverse orange brown band broadly
interrupted medially; disc blackish brown. Abdomen
blackish brown, slightly lighter ventrally, connexiva
margined with orange brown. Antennae brown; ante-
rior femora yellowish, distally darkened, tibia brown;
middle, hind legs dark brown to piceous, femora ba-
sally lighter; coxae, trochanters luteous, tinged with
fuscous.
Structural characteristics. — Apterous male: head of
moderate length, declivant anteriorly, with weak im-
pressed median line; length 0.33; width of eye/inter-
ocular space, 0.14/0.28. Pronotum long, covering
metanotum, thickly set with obscure foveae, humeri
depressed; length:width, 0.37 : 0.81. Metanotum
length:width, 0.28 : 0.79. Abdominal tergites not
shining; tergites II-VI subequal in length (0.12-0.16),
VII longer (0.30).
Abdominal venter set with short appressed setae;
venter V posteriorly set with two (1+1) tufts of dense
dark erect setae on either side of midline; venter VI
with caudal margin medially produced, set with a
large quadrate tuft of dense dark erect setae; venter
VII excavated medially along posterior margin. Legs,
antennae thickly clothed with short to moderate
length setae, with scattered longer setae. Middle and
hind legs unarmed; fore femur lacking denticles or
spinules; fore tibia with grasping comb extending 1/2
the length of the tibia.
Antennal formula I: II: III: IV; 0.30: 0.21 : 0.28
: 0.44.
Proportions of legs as follows: Femur, tibia, tarsal
1, tarsal 2 of fore leg, 0.51 : 0.47 : 0.21 : 0.0; of mid-
dle leg, 0.70 : 0.63 : 0.12 : 0.23; of hind leg, 0.74 :
0.93 : 0.16 : 0.28.
Proctiger produced anteriorly into a bifurcate pro-
cess (fig. 21). Parameres vestigial or absent.
Apterous female: Similar to male in most respects,
72
but somewhat larger; abdomen narrowing posterior-
ly. Connexiva vertical, touching at caudal extreme;
caudally rounded; thickly set along posterior margin
with long stout dark setae; tuft of stiff posterolateral-
ly directed setae near connexival margin of lateroster-
nite VII of females pronounced, more so than in oth-
er species.
Macropterous male: Similar to apterous male in
many respects, but broader, humeri more pro-
nounced. Hemelytra always with light streaks in each
of the two basal cells, often with light areas in all
closed cells.
Macropterous female: Similar to apterous female in
many respects, but broader, humeri more pro-
nounced. Hemelytra with light areas in each of the
four basal closed cells.
Remarks
Comparative notes. — Easily recognized by the bi-
furcate process on the male proctiger (fig. 21) and the
laterally compressed, boat-shaped abdomen in fema-
les. This species is similar in size to N. gladius sp. n.,
and larger than N. cyclops sp. n., but smaller than eit-
her N. compactus sp. n. or N. sepik sp. n.
Biological notes. — See discussion under N. gladi-
us Sp. n.
Etymology. — The name ‘rabire, a noun in apposi-
tion, refers to the Nabire River type locality.
Distribution. — Northern New Guinea (fig. 30).
Neusterinsifer sepik sp. n.
(figs. 20, 24, 30)
Type material: Holotype, apterous male: Papua
New Guinea, East Sepik Prov., Yemogu Creek, 2 km
Wa iliring 12 Sep 41985 CIS OS 7]. ED
Polhemus (BPBM). - Paratypes: 11 apterous males, 6
apterous females, same data as holotype (JTPC); 1 ap-
terous male, 4 apterous females, 1 macropterous fe-
male, East Sepik Prov., Mandi Creek, E. of Wewak,
10 Sept. 1983, CL 1797, J. T. & D. A. Polhemus (-
JTPC); 4 apterous males, 1 apterous female, East Sepik
Prov., Brandi Creek, nr. Wewak, 10 Sept. 1983, CL
1796, J. T. & D. A. Polhemus (JTPC).
Diagnosis
Size. — Apterous male, length 2.27-2.41 mm (x =
2.35, n = 10); width 0.94-1.01 mm (x = 0.99, n =
10). Apterous female, length 2.37-2.63 mm (x =
2.48, n = 10); width 0.97-1.15 mm, (x = 1.04, n =
10). Macropterous female, length, x = 2.62, n = 1);
width, x= 1.15, n = 1)
Colour. — Apterous male: Ground colour blackish
brown, venter slightly lighter, entire dorsum and la-
POLHEMUS & POLHEMUS: Microveliinae from New Guinea
Fig. 29. Distribution of
species of Tarsovelia gen. n.
— Square = Tarsovelia dani
sp. n.; Circles = Tarsovelia
alta sp. n.; Triangle =
Tarsovelia arfak sp. n.
Figure 30. Distribution of
species of Neusterinsifer
gen. n. — Square =
Neusterinsifer sepik sp. n.;
Circles = Neusterinsifer
compactus sp. n.; Triangle =
Neusterinsifer cyclops sp. n.;
Diamond = Neusterinsifer
nabire sp. n.; Half filled
square = Neusterinsifer gla-
dius sp. n.
terotergites covered with fine appressed golden pu-
bescence; entire body except thoracic venter covered
with long erect setae. Head black, orange brown be-
neath; rostrum fuscous medially, piceous distally.
Pronotum with anterior transverse orange brown
band barely interrupted medially; disc blackish
brown. Abdomen blackish brown, lighter ventrally,
connexiva margined with orange brown. Antennae
brown; anterior legs yellowish, distally darkened;
middle, hind legs dark brown to piceous dorsally,
ventrally yellowish, femora basally lighter; coxae, tro-
chanters luteous, tinged with fuscous.
Structural characteristics. — Apterous male: head of
moderate length, with weak impressed median line;
length 0.27; width of eye/interocular space,
0.14/0.30. Pronotum long, covering metanotum,
thickly set with obscure foveae, humeri depressed;
length:width, 0.37 : 0.95. Metanotum length:width,
0.28 : 0.95. Abdominal tergites not shining; tergites
II-VI subequal in length (0.14-0.16), VII longer
(0.30).
Abdominal venter set with short appressed setae;
venter VI posteriorly set with two (1+1) tufts of dense
dark erect setae on either side of midline, with hair
free sulcus between; venter VII broadly depressed me-
dially, weakly carinate along posterior margin. Legs,
antennae thickly clothed with short to moderate
length setae, with scattered longer setae. Middle and
hind legs unarmed; fore femur bearing an elongate
patch of stiff erect black spinules basally; fore tibia
with a swelling distally bearing 5-7 erect short stiff se-
tae (fig. 24).
Antennal formula I: II : III: IV; 0.33 : 0.26 : 0.37
: 0.47.
Proportions of legs as follows: femur, tibia, tarsal 1,
tarsal 2 of fore leg, 0.65 : 0.53 : 0.28 : 0.0; of middle
leg, 0.84 : 0.79 : 0.16 : 0.28; of hind leg, 0.93 : 1.07
019028: ’
Proctiger produced anteriorly into a long broad
sinuate ensiform process (fig. 20). Parameres vestigial
or absent.
Apterous female: Similar to male in most respects,
but somewhat larger; connexiva vertical; caudal ex-
treme forming an acute angle.
Macropterous female: Similar to apterous female in
many respects, but broader, humeri more pro-
nounced. Hemelytra with light areas in each of the
four basal closed cells.
75
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Remarks
Comparative notes. — Recognized by the large,
elongate process on the male proctiger that comes to
a rounded point (fig. 20), and the elongate patch of
black denticles on the inner surface of the male fore-
leg (fig. 24).
Biological notes. — Yemogu Creek at the type local-
ity was a slow flowing stream running through a rock
bed, and containing numerous large pools shaded by
slightly disturbed primary rain forest. N. sepik was
found here in sheltered areas along the margins of the
stream pools, and in similar habitats at Brandi River
and Mandi Creek, two lowland rainforest streams en-
tering the sea east of Wewak.
Etymology. — The name sepik, a noun in apposi-
tion, refers to the district of Papua New Guinea from
which the type series was taken.
Distribution. — Northern New Guinea (fig. 30).
ACKNOWLEDGEMENTS
We are pleased to acknowledge the invaluable assistance
of the following persons who aided us with permits, guidan-
ce, transportation and logistics, and without whose help and
patience our field surveys in Indonesia could not have been
accomplished: Drs. Rosichon Ubaidillah, Dr. Soetikno
Wiroatmodjo, Dr. Mohammed Amir, and other personnel
from the Indonesian Institute of Sciences (LIPI), Bogor; Mrs.
Moertini, Mr. Sanchoyo, the other staff members of the LIPI
office in Jakarta; Mr. Benny Lesomar, Natrabu Corp., Biak;
Mr. Jacob Bakabessy and Mr. Bernardus Sambery, PHPA,
Manokwari; and Mr. Henk van Mastrigt, Jayapura.
Special thanks are also due to the following persons who
aided us with field surveys in Papua New Guinea: Stan and
74
Kris Niemi, Mt. Hagen; and especially John Ismay, former-
ly of the Bureau of Primary Industry, Konedobu, and now at
the Hope Museum of Entomology, Oxford.
Holotypes of all new species described herein are deposit-
ed in the Bishop Museum, Honolulu (BPBM), or the U.S.
National Museum of Natural History, Washington, D. C. (-
USNM); paratypes are held in the J. T. Polhemus collection,
Englewood, Colorado (JTPC), and where numbers permit
will be distributed to the institutions noted above as well as
the Museum Zoologicum Bogoriense, Bogor, Indonesia
(MZB), and the Universitets Zoologisk Museum,
Copenhagen (ZMUC). We also thank Dr. Martin Baehr of
the Zoogische Sammlung des Bayersichen Staates, Munich
(ZBSM) for allowing us to examine specimens held under his
care.
This research was sponsored by a series of grants (2698-
83, 3053-85, 4537-91) from the National Geographic
Society, Washington, D.C., and by grant BSR-9020442
from the National Science Foundation, Washington, D.C.
We thank these organizations for their continued support
on research into the systematics and zoogeography of aquat-
ic Heteroptera.
REFERENCES
Andersen, N. M. 1982. The semiaquatic bugs (Hemiptera,
Gerromorpha). Phylogeny, adaptations, biogeography
and classification. — Scandinavian Science Press,
Klampenborg, Denmark, Entomonograph 3, 455 pp.
Andersen, N. M. 1989. The Old World Microveliinae
(Hemiptera: Veliidae). II. Three new species of Baptista
Distant and a new genus from the Oriental region. —
Entomologica Scandinavica , 19: 363-380.
Received: 19 November 1993
Accepted: 25 March 1994
R. ROZKOSNY! & D. KOVAC?
' Masaryk University, Brno? Forschungsinstitut Senckenberg, Frankfurt
ADULTS AND LARVAE OF TWO PTECTICUS LOEW
FROM PENINSULAR MALAYSIA (DIPTERA,
STRATIOMYIDAE)
Rozkosny, R. & D. Kovac, 1994. Adults and larvae of two Ptecticus Loew from Peninsular
Malaysia, (Diptera, Stratiomyidae). — Tijdschrift voor Entomologie 137: 75 - 86, figs. 1-43.
[ISSN 0040-7496]. Published 15 July 1994.
Descriptions and redescriptions of larvae and adults of P. malayensis sp. n. and P. longipennis
(Wiedemann) are given, based on material from Malaysia. The phylogenetic relationships of
the two species and the systematic position of the genus based on larval and adult characters are
briefly discussed, and biological notes on larvae are provided. The diagnostic characters of the
larvae and imagines, including the male genitalia, are illustrated. A generic key to the larvae of
the subfamily Sarginae is given.
Correspondence: R. Rozkoënÿ, Department of Environmental Studies, Faculty of Science,
Masaryk University, Kotläïskä 2, 611 37 Brno, Czech Republic.
Key words. — Peninsular Malaysia; Stratiomyidae; larvae, key; Ptecticus new species, life histo-
ry.
The genus Precticus Loew belongs to the subfamily
Sarginae and differs in the adult stage from the close-
ly related genus Sargus Fabricius by the distinctly
prominent pedicel on the inner side of the antenna.
According to the recent catalogue by James (1975),
31 valid species have been described from the
Oriental region but many of them are poorly known.
The male genitalia as the most important criterion for
species discrimination, for example, have not yet been
examined in most Oriental species and the taxonom-
ic status of many species needs a re-evaluation.
The junior author, while studying insects on bam-
boo (Gigantochloa scortechinii Gamble) in Peninsular
Malaysia, found larvae of Stratiomyidae living inside
bamboo internodes. Following our study on
Camptopteromyia fractipennis de Meijere (Rozkosny
& Kovac 1991) the present paper represents the sec-
ond modern contribution to the taxonomy, morphol-
ogy and biology of the Oriental Stratiomyidae.
Two species are treated here, both reared from larvae
found under semi-natural conditions. The first of them,
P. longipennis (Wiedemann), is a well distinguishable
species, but we decided on re-description in order to in-
clude important diagnostic characters not mentioned in
the original description, including the male genitalia.
The second species is described here as new; it differs
from all known Oriental Ptecticus species. Larvae and
puparia of both species are described for the first time,
and their descriptions enable us to propose generic
characters for Ptecticus in the larval stage.
Ptecticus longipennis (Wiedemann, 1824)
(figs. 1-10, 26-32)
Diagnosis
An easily recognizable, chiefly blue black and yel-
low species from the cingulatus-group with conspi-
cuously long, and apically darkened wings.
Measurements: Body 8.2-11.5 mm, wing 10.8-
13.0 mm.
Male. — Head hemispherical, deeply concave poste-
riorly, bare eyes touching on frons (figs. 1-2). Upper
part of frons elongate, triangular, subshining black and
sparsely covered with fairly long brownish hairs.
Ocellar triangle relatively narrow and low, barely vis-
ible in lateral aspect (fig. 2). Lower frons and face pro-
tuberant, white to pale yellow, white frontal callus slen-
der, nose-shaped. Face with dense yellow hairs.
Antennae yellow, arista brown, both basal segments
yellow haired. Pedicel with prominent projection on
inner side. Flagellum wider than long, about as long as
pedicel. Postocular margin with a complete fringe of
erect hairs. Proboscis pale yellow, somewhat swollen
but relatively slender, with two-segmented labellae.
Thorax bright yellow in ground colour, with exten-
sive dark pattern. Mesonotum chiefly shining bluish-
black, leaving large anterior corners and postalar calli
yellow. Scutellum dark, with margin broadly brown-
ish or yellow. Pleura yellow, with large brownish to
bluish-black spots on greater part of anepisternum,
75
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 1-6. Precticus longipennis, adult characters. — 1, male head, dorsal view; 2, the same, lateral view; 3, female head, dorsal
view; 4, dorsal part of male terminalia; 5, aedeagal complex; 6, ventral part of male terminalia.
lower half of katepisternum and anterior parts of ane-
pimeron and katepimeron. Laterotergite shining
brown and mediotergite behind scutellum complete-
ly bluish-black. Narrow subnotopleural stripe, upper
margin of katepisternum, and, especially, propleuron
pale, often whitish yellow.
Wings slightly greyish infuscate, with yellow pte-
rostigma and darkened apical part reaching almost
half way between apex and discal cell. Dense micro-
trichia absent in wing base, alula, subcostal cell and
greater part of posterior cubital cell. Anterior cross-
vein situated just at beginning of R,,, M, shortened,
not reaching wing margin. Narrow squamae white to
yellow, pale yellow haired. Halteres yellow, with
darkened knob.
Legs including coxae chiefly yellow, only hind pair
partly darkened. All femora distinctly thickened in
basal half. Hind coxa and femur yellow as in fore and
middle legs, hind tibia and narrow base of basitarsus
dark brown to black. Rest of basitarsus and all follow-
ing tarsal segments white, with whitish hairs at least
on dorsal side.
76
Abdomen shining bluish-brown to black, with yel-
low transverse stripes and pleural membrane.
Transverse stripes broad on the posterior, and narrow
on the anterior margin of each tergum of segments 2-
4, apical terga completely black. Transverse stripes
sometimes reduced to broad yellow sidemarkings on
posterior corners of segments. Male genitalia (figs. 4-
6) rotated by 90° along longitudinal axis, so that
epandrium is in right lateral position. Apart from
this, hypopygium symmetrical. Epandrium almost
round and compact, tightly connected with para-
procts posteroventrally. Epiproct of proctiger narrow,
bearing slender, rodlike cerci. Synsternum deeply
emarginate, with slender and bipartite medial process
shielding short, symmetrical, partly reduced aedeagal
complex from ventral side.
Female. — Eyes separated by shining black frontal
stripe being anteriorly somewhat broader than ocellar
triangle (fig. 3). White frontal callus bulbose, wider
than in male. Scutellum dark, its posterior margin
slightly yellowish. Dark pleural spots as in male but
partly reduced on katepisternum. Abdomen shining
bluish black, with transverse yellow stripes at posteri-
or margins of terga 2-4 extending to anterior margins
of following terga. Yellow stripes about half as wide as
dark stripes between them, somewhat broader than in
male. Dark pattern of hind legs as in male. Cerci with
two segments, slender and relatively long.
Larva (and puparium). — Subcylindrical, with pro-
minent head, slightly arched lateral margins of body
segments and distinctly angulate though generally
Lb
4
ROZKOSNY & KOVAC: Ptecticus
rounded anal segment. Cuticle with usual mosaic ap-
pearance, some cells somewhat larger and darker,
forming almost complete double rows at boundary of
body segments and groups in sublateral position.
General colour yellowish brown, with diffuse brown
pattern consisting of broad, longitudinal, undulate
bands. Anal segment yellow, with large anterodorsal
brownish spot. Ventral side paler, especially in me-
dian third where brownish spots are almost indistinct.
Head (figs. 7-8, 26-27) moderately flat, subconical.
Vestiges of mandibular-maxillary complexes about as
Figs. 7-10. Ptecticus longipennis, larval characters. — 7, head and first thoracic segment, dorsal view (A, antenna; Ad, antero-
dorsal setae; Cf, clypeofrontal setae; D, dorsal setae; DI, dorsolateral setae; E, eye prominence; L, lateral setae; Lb, labral se-
tae; VL3, posterior ventrolateral setae); 8, the same, ventral view (SI, sublabral setae; V, ventral setae; VL1-3, ventrolateral se-
tae; V3, posterior ventral setae); 9, last abdominal segments, dorsal view ( Sa, subapical setae); 10, the same, ventral view (Ap,
apical setae; as, anal slit; L1 + L2, lateral setae; sp, sternal patch).
VY.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs 11-19. Ptecticus malayensis sp. n., adult characters. — 11, male head, dorsal view; 12, the same, lateral view; 13, mesono-
tum pattern, dorsal view; 14, male terminalia, lateral view; 15, female abdomen, dorsal view; 16, antenna, inner side; 17, dor-
sal part of male terminalia; 18, aedeagal complex; 19, ventral part of male terminalia.
78
long as labrum in dorsal view, low and rounded an-
tennae placed anteriorly, their apical segment about
as high as broad at base. Eyes relatively little promi-
nent, posterior part of head with two rounded protu-
berances on each side.
Three thoracic segments about same shape as ab-
dominal segments 1-7, first thoracic segment with
lateral vestiges of prothoracic spiracles. Thoracic seg-
ment 3 and abdominal segments 1-7 with small,
round, dorsolateral, probably non-functional larval
spiracles. In puparia, abdominal segments 2-5 with
very inconspicuous, short, rodlike pupal respiratory
horns near lateral margin. Very slender and long mid-
sternal patch on abdominal segment 6 consisting of
112-114 pale cells (figs. 10, 31). Rounded and much
smaller sternal patch with 18-19 cells usually at poste-
rior margin of abdominal segment 5 (fig. 32). Anal
segment (figs. 9-10, 28) distinctly emarginate
between insertions of marginal setae, with a pair of
low protuberances distally. Opening of spiracular
chamber with undulate posterior margin and short
pennate setae (figs. 29-30). Anal slit on ventral side
with somewhat swollen marginal lips, boundered
proximally by usual transverse fold. Two low sym-
metrical protuberances developed around insertions
of posterior ventral setae.
General scheme of chaetotaxy as in other known
genera of Sarginae (cf. Rozkosny 1982). All setae on
head short, labral setae not longer than others, spine-
like sublabral setae well developed. Anterior clypeo-
frontal setae inserted almost at level of eye promi-
nences. Setae on thoracic and abdominal segments
arranged in transverse rows (3 pairs of dorsal and 2-3
pairs of ventral setae) and lateral groups (2-4 setae).
Thoracic segment 1, on each half, with 2 anterodor-
sal, 3 dorsal, 1 dorsolateral and 1 ventrolateral seta.
All 3 thoracic segments only with 2 pairs of ventral se-
tae. Lateral group of setae on thoracic segments con-
sisting of 2 setae only (1 dorsolateral and 1 ventrolat-
eral).
On lateral side of abdominal segments, 4 setae (1
dorsolateral, 2 lateral and 1 ventrolateral) on abdom-
inal segments | and 6-7, but only 2 setae (probably |
ventrolateral and | lateral) on abdominal segments 2-
5. All marginal setae on anal segment of about same
length, virtually not longer than setae on preceding
abdominal segments. Five pairs of ventral setae on
anal segment well visible, two distal pairs inserted
closely together (fig. 10). Especially dorsal and ventral
setae on body segments distinctly pubescent. Length
9.3-10.6 mm, maximum width 2.6-3.0 mm (based
on 4 puparial exuviae).
Material examined. — Peninsular Malaysia, University of
Malaya Field Studies Centre, Ulu Gombak, Selangor
(3°20’N, 101°45’E, altitude 250 m), 1d from 10 May, 2d
ROZKOSNY & KOVAC: Ptecticus
from 12 May and 19 from 16 June 1991, all specimens ex
puparia, D. Kovac leg. Material deposited in the
Forschungsinstitut Senckenberg, Frankfurt (SMFD) (28),
and in the Faculty of Science, Masaryk University, Brno
(MUB) (16 and 19).
Remarks
Distribution. — India, Malaya, Sumatra, Java,
Borneo, Philippines.
Discussion. — According to Brunetti (1923) this
slender species with long legs and wings belongs to
the cingulatus species group. It differs from the other
species of this group by the darkened wing tip, a diag-
nostic character shared only with P. sumatranus
Enderlein. The latter species, however, possesses the
normal, not elongate wings and its mesonotum is red-
dish yellow, not dark with a metallic shine. Of the few
Ptecticus species in which the male genitalia have been
illustrated, only P. okinawensis Ouchi may belong to
the same group, as shown by the compact epandrium
and deeply emarginate synsternum (cf. Nagatomi
1575)
Ptecticus malayensis sp.n.
(figs. 11-25, 33-39)
Type material. — Holotype, male: Peninsular Malaysia,
University of Malaya Field Studies Centre, Ulu Gombak,
Selangor (3°20’N, 101°45’E, altitude 250 m), ex puparium
15 November 1991, D. Kovac leg., SMFD. — Paratypes 3d
29: Same locality, 38 reared 15 November, 12 23
November, 1 2 20 December 1991; 16 19 SMFD, 26 19
MUB.
Diagnosis
A species of the cingulatus group with longitudinal-
ly striped mesonotum and a conspicuously spindle-
shaped abdomen in both sexes.
Measurements: Body 11.2-12.5 mm, wing 11.8-
12.2 mm.
Male (holotype). — Head hemispherical, with con-
spicuously protuberant white frontal callus (figs. 11-
12). Eyes separated by yellow, sparsely black-haired
frontal stripe being in its narrowest point about as
wide as distance between outer margins of posterior
ocelli. Postocular area narrowly protuberant in dorsal
view. Postocular hairs fine and moderately long, in
several rows. Antennae ochrous yellow, scape and
pedicel black-haired. Pedicel with elongate process on
inner side (fig. 16), annulation of flagellum barely vis-
ible. Arista in subapical position, bare. Bi-partite la-
bellae of proboscis distinctly swollen, white in ground
colour and with short white hairs. Posterior part of
head deeply concave.
Thorax yellow in ground colour, with extensive
79
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 20-25. Ptecticus malayensis sp.n., larval characters. — 20, head and first thoracic segment, dorsal view; 21, the same, ven-
tral view; 22, last abdominal segments, dorsal view; 23, the same, ventral view; 24, scheme of setae on lateral wall of 6th ab-
dominal segment (DI, dorsolateral seta; st, larval spiracle; VI, ventrolateral seta); 25, scheme of setae on lateral wall of abdom-
inal segment 3 (ph, pupal respiratory horn; st, larval spiracle).
brown to blackish pattern. Mesonotum (fig. 13) with to black spots in central part of anepisternum, anteri-
3 broad black stripes reaching its posterior margin. or side and posterior corner of katepisternum, poste-
Pleura predominantly yellow with several brownish rior half of katepimeron and anterior third of latero-
80
tergite. Also posterior margin of mediotergite broadly
brownish. Thoracic pubescence short and mostly ad-
pressed, yellow on pleura and predominantly brown
to black on mesonotum and scutellum.
Wings shaded with grey, veins along anterior mar-
gin including discal cell yellow, other veins brown ex-
cept for bases. Anterior cross-vein placed at basis of
radial fork R,, M, not reaching wing margin. Dense
microtrichia covering virtually the whole wing mem-
brane, only wing base, alula and posterior cubital cell
mostly bare. Squamae very narrow and inconspicu-
ous, with brownish margin and yellow hairs. Halteres
with yellow stem and blackish knob.
Legs yellow with extensive blackish pattern on hind
pair. Both anterior pairs mainly yellow, including
coxae, only apical half of mid-femora somewhat
brownish. Hind coxae brownish on anterior side,
hind femur mainly dark except for both ends. Also
hind tibia chiefly brown but this darkening more pro-
nounced along dorsal side and in mid-ring. Hind tar-
si black but basal two thirds of basitarsus and last tar-
sal segment pale yellow.
Abdomen spindle-shaped, broadest behind middle,
yellow with brownish black transverse bands on terga
(fig. 15). First tergum yellow with a transverse brown
spot in middle and following terga with relatively
broad posterior margin and narrow anterior margin
yellow. Venter yellow but more brownish in apical
half. Genitalia (figs. 14, 17-19) brown, distinctly
asymmetrical, although dorsal part only slightly so.
Cerci pointed, with tips oriented ventrally (figs. 14,
17). Synsternum strongly asymmetrical in proximal
half, dististyli almost hoodlike in ventral view (fig.
19). Aedeagal complex almost S-shaped in lateral as-
pect (fig. 14), right paramere conspicuously reduced
(fig. 18).
Female. — As male but with broader wings and
somewhat wider abdomen in apical half. Frontal
stripe only slightly broader than in male, particularly
in anterior half, white frontal callus distinctly bipar-
tite and lower, antennae relatively larger. Colours of
thorax, legs and abdomen virtually the same, only
darkening of hind legs more extensive, leaving only
bases of femora and knees yellow. Hind tarsi coloured
as in male. Terminalia dark brown, cerci with two
narrow black segments (fig. 15).
Larva (and puparium). — Larvae on average stouter
and longer than those of P. longipennis, often pre-
dominantly reddish brown, though dark pattern con-
sisting of dark undulate longitudinal bands, especial-
ly in puparia, mostly distinct. Integument rough and
sclerotised, with usual mosaic surface structure. Some
cuticular cells forming inconspicuous pattern as in
preceding species. Head (figs. 20-21, 33-34) promi-
ROZKOSNY & KOVAC: Precticus
nent, strongly sclerotised, partly retracted into first
thoracic segment, distinctly longer than wide and
conspicuously flattened, rather subtriangular in dor-
sal view. Antennae very small and low, distinctly clos-
er to eye prominences than in P. longipennis, apical
segment barely prominent. Eyes conspicuously pro-
tuberant, posterolateral tubercles less distinct, not
doubled as in P. longipennis.
Tracheal system as in larva of P. longipennis, in-
cluding vestiges of larval spiracles and very small lat-
eral pupal respiratory horns on abdominal segments
2-5. Sternal patch in middle of abdominal segment 6
drop-like to elongate oval, consisting of 90-116 cells
(fig 23). Similar but much smaller and rounded ster-
nal patch at posterior margin of sternum 5 formed by
17-20 cells. Anal segment (figs. 22-23, 35) almost
round posteriorly, with a small incision in middle of
distal margin. Opening of posterior spiracular cham-
ber with arched and finely fringed posterior margin (-
figs. 36-37). Anal slit distinctly emarginate, some-
what longer than one third of anal segment.
General scheme of chaetotaxy as in P. longipennis
although setae relatively shorter and more pubescent,
especially in marginal setae of anal segment. Anterior
clypeofrontal setae inserted almost between eye
prominence and tip of labrum (fig. 20). Configura-
tion of setae on lateral wall of abdominal segments
and ventral side of anal segment identical with those
of preceding species and probably confirming generic
characters in this way: 4 setae on lateral wall of ab-
dominal segments | and 6-7, 2 setae on abdominal
segments 2-5 (figs. 24-25) and 5 pairs of ventral setae
on anal segment present (fig. 23). Length 10.0-13.4
mm, maximum width 3.6-5.0 mm (based on 2 larvae,
5 puparia and 3 puparial exuviae).
Discussion
The new species presumably belongs to the cingu-
latus species group as defined by Brunetti (1923).
Three species of this group have dark longitudinal
stripes on the thorax: P. brevipennis Rondani, P. in-
tensivus Brunetti and P. kambangensis de Meijere. The
first was redescribed by van der Wulp (1885) and
Brunetti (1923). It is, in contrast to P. malayensis sp.
n., characterized by the shining blue black frontal stri-
pe and by much narrower and shorter stripes on the
mesonotum, the black tibiae and bases of the basitar-
si of the hind leg. P. intensivus is, according to the ori-
ginal description by Brunetti (1927), closely related
to P. brevipennis and like that species should have a
black frontal stripe. Moreover, it also differs in the co-
loration of the leg (the fore tarsi are partly black, only
the basal third of the hind femur is darkened and the
hind tarsi are completely black). P. kambangensis dis-
tinctly differs by the pleura being shining blackish
81
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
82
brown with a yellow band, and by the hind basitarsus,
which is completely whitish as is the basis of the fol-
lowing tarsal segment.
From the other Oriental Ptecticus species only P.
tricolor van der Wulp displays the longitudinally
striped mesonotum. Brunetti (1923) considered this
species to be an intermediate form with uncertain re-
lationships. According to the original description and
illustration (van der Wulp in de Meijere 1904) P. tri-
color differs from the new species by the colouring of
the legs (hind tibia and tarsi completely black) and
abdomen (segments 2-4 entirely red).
The colour of the leg and partly also the male gen-
italia of the new species resemble the East Palaearctic
P. sinchangensis Ouchi redescribed and figured re-
cently by Nagatomi (1975). However, this species
does not have a striped mesonotum, and its male gen-
italia, including the well developed aedeagal complex,
are symmetrical.
BIOLOGICAL NOTES
The larvae of Ptecticus longipennis were found in
felled bamboo culms whose internodes — now lying in
the horizontal position on the ground — had been
sawn in such a way that the upper half formed a hin-
ged lid that could easily be opened for inspection and
than closed again. Two lateral slits, about 2 mm wide,
provided access to the interior. The internodes were
partly filled with water, and the larvae stayed in the
moist part above the water surface (fig. 43). They ap-
peared about two months after the culms had been
felled. During the six-months study period larvae of
Ptecticus longipennis were found only in three out of
over one hundred internodes prepared in the descri-
bed manner; moreover, these three internodes were
adjacent to each other, i.e. the eggs were probably laid
by the same female. This indicates that bamboo in-
ternodes are only an occasional habitat for the larvae
of Precticus longipennis.
Ptecticus malayensis, in contrast, was found in bam-
boo more frequently. The larvae of this species were
restricted to bamboo shoots that had died recently
(fig. 40). Judging from the traces of insect attack
found in the internodes, death of the shoots was
probably caused by a weevil, Cyrtotrachelus sp.
(Coleoptera, Curculionidae), adults of which had
been observed on bamboo shoots. The dead shoots
were 2.5 m to 6 m high. At the time of felling the
lower internodes were still green while the upper part
ROZKOSNY & Kovac: Precticus
had turned dark brown and begun to decay. In the
lower part the rotten section (c. 1 m from the top) in
some cases a small hole (diameter c. 1-2 mm) was
found which led into the interior of the culm.
Beginning at this hole a groove in the inner wall led
upwards, piercing the node septa (usually c. 10-15)
and gradually becoming wider and deeper (fig. 42).
At several points along this depression there were
holes through the internode wall to the outside which
were covered by the culm sheets and therefore hidden
from outside view. Just below the shoot tip there was
no segmentation into internodes. In this section,
which was moist and gave off a putrid smell, the culm
sheets had been penetrated with a circular opening c.
1.5 cm in diameter. In one case 2 grooves and circu-
lar openings were found in the same shoot.
Apparently the Cyrtotrachelus larva had worked it-
self up all the way to the nutritious apical meristem
from the small entrance hole bored by the female,
feeding and grooving along the way. After feasting on
the apical meristem the fully grown larva then prob-
ably broke to the outside tu pupate in the soil, as de-
scribed by Witt (in Beeson 1941: 209) for
Cyrtotrachelus longipes F.
Ptecticus malayensis probably uses a fresh Cyrtotra-
chelus emergence hole to enter the bamboo shoot.
Although all shoots attacked by Cyrtotrachelus died,
there was probably still some limited growth after de-
parture of the weevils as indicated by a shift of the
holes in the different culm sheets relative to each oth-
er (fig. 41). Consequently, at the time of felling there
was no longer a contiguous opening all the way to the
interior of the culm. The larvae of Ptecticus malyensis
were found in all internodes connected by holes. In
the lower internodes with punctured septa water usu-
ally accumulated, and dead stratiomyid larvae were
found there on several occasions. Pupation of Ptecti-
cus malayensis took place in the upper section of the
shoot, behind the Cyrtotrachelus escape hole (fig. 41).
Probably the freshly eclosed flies had to sqeeze them-
selves past the sheets to escape from the bamboo. The
adult flies left before the rotten shoot tips fell to the
ground. Five out of twelve investigated bamboo
shoots were occupied by Ptecticus malayensis, with a
maximum number of 22 larvae per shoot. In two
shoots larvae of Solva completa de Meijere (Xylomyi-
dae, Diptera) were found together with Ptecticus ma-
layensis.
Only a few literature accounts on larvae of Precticus
are available. Lindner (1928), who superficially de-
Figs. 26-32. Ptecticus longipennis, larval structures (SEM-photographs). — 26, head and fore margin of first thoracic segment,
dorsal view; 27, the same, lateral view; 28, last abdominal segments, ventral view; 29, opening to spiracular chamber, poste-
rior view; 30, short pennate setae on ventral lip of spiracular chamber opening; 31, sternal patch on abdominal segment 6; 32,
sternal patch on abdominal segment 5.
83
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 33-39. Ptecticus malayensis sp.n., larval structures (SEM-photographs). — 33, head and fore margin of first thoracic seg-
ment, dorsal view; 34, head and two thoracic segments in lateral view; 35, last abdominal segments, ventral view; 36, open-
ing to spiracular chamber, posterior view; 37, pennate setae on lips of spiracular chamber opening; 38, sternal patches on ab-
dominal segment 6; 39, sternal patch on abdominal segment 5.
84
ROZKOSNY & Kovac: Ptecticus
Figs. 40-43. Habitat of the larvae of Ptecticus malyensis sp.n. and P. longipennis. — 40, dead shoot tip of Gigantochloa scortechi-
nit. The arrow points to the emergence hole of the weevil larva, Cyrtotrachelus sp., which was probably used by P. malayensis
to enter the hollow interior of the shoot; 41, emergence hole of Cyrtotrachelus sp. after removal of several culm sheets. Two
puparia of P. malayensis are visible (arrows). As can be seen, the openings of the culm sheets have shifted relative to each oth-
er; 42, the hollow interior of the shoot. Arrow points to a groove made by the larva of Cyrtotrachelus sp.; 43, larva of Ptecticus
longipennis on the moist interior wall of an internode laying on the ground.
scribed a larva of P. testaceus Fabricius from South
America, pointed out that larvae were found in decay-
ing cactuses, oranges and lemons. Engel & Cuthbertson
(1939), who examined a larva of African P. posticus
Wiedemann, found the larvae in the semi-liquid mass
of decaying Conopharyngia johnstoni Stapf. (Apocyna-
ceae) fruits together with other dipterous larvae.
Larvae of North-American Ptecticus trivittatus
(Say) were collected from a corn compost pile, decay-
ing tomatoes, garbage and fruit, from watermelon
rinds, rotting paper and a fungus (McFadden 1967).
Our observations suggest that P. longipennis and P.
malayensis are unspecialized phytosaprophages. These
findings conform to the previously reported larval
mode of life within the subfamily Sarginae: sapropha-
gy with some tendency to coprophagy as known in
many European species.
RELATIONSHIPS OF Ptecticus LOEW
Only one reliable character separates the genus
Ptecticus Loew from the related Sargus Fabricius, viz.
the shape of the pedicel projecting into the elongate
process on the inner side of the antenna. However,
this character may be indistinct in some species, and
its generic relevance therefore seems disputable. A
comparison of the structure of the male genitalia is
not possible for all species of the two genera because
it has not been included in all descriptions. It may ne-
vertheless be stated that the differences found among
some Ptecticus species seem to be more pronounced
than those among the European genera of the
Sarginae. This fact indicates the necessity of a revision
of the generic concept of Ptecticus.
Judging from the description of the larva of North-
American P. trivittatus (Say) by McFadden (1967)
and the descriptions of the two larvae presented here,
Ptecticus appears, on the other hand, well separated
from the genera of Sarginae in the larval stage. First of
all, in addition to a long and slender sternal patch on
abdominal segment 6, a small and rounded sternal
patch on abdominal segment 5 is developed.
Relatively short setae on the head as well as on the
anal segment, the configuration of setae on the lateral
wall of abdominal segments and the presence of five
pairs of ventral setae on the anal segment are cosid-
85
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
ered diagnostic generic characters. So, the known lar-
vae of Sarginae may be distinguished according to the
following idenufication key:
1. Setae on anal segment shorter than length of pe-
DUITIMATE SÉDIMENTS een D
— At least some setae on anal segment longer than
penultinnatelse nnen Ge 4
2. Eye prominence anterior to protruding lateral tu-
bercle (according to McFadden 1967) ................
AIA A OE RU ee Merosargus Loew
— Eye prominence on protruding lateral tubercle
3. Abdominal segment 5 without sternal patch, four
pairs of ventral setae on anal segment, 2-4 setae
on lateral wall of abdominal segment 1, 6-7, and
always one seta less or equal number of setae on
abdominal segments 2-5 ........... Sargus Fabricius
— Abdominal segment 5 with a small but distinct
sternal patch in addition to sternal patch on seg-
ment 6, five pairs of ventral setae on anal seg-
ment, four setae on lateral wall of abdominal seg-
ments |, 6-7, and only two setae on abdominal
segments 2-5 (i.e. two setae less) … Ptecticus Loew
4. Anterior labral setae short, apical lobes on anal
segment prominent ............ Chloromyia Duncan
— Anterior labral setae strikingly long, apical lobes
on anal segment less distinct .. Microchrysa Loew
ACKNOWLEDGEMENTS
Thanks are due to K. Fiedler and R. Klein for re-
viewing the manuscript. This work is part of a re-
search project funded by the Deutsche Forschungsge-
meinschaft and supported by the Socio-Economic
Research Unit of the Government of Malaysia and
Universiti Malaya. Special thanks are due to I. Azarae
for supplying the facilities of the Ulu Gombak Field
Studies Centre.
86
REFERENCES
Beeson, C.F.C., 1941. The ecology and control of the forest
insects of India and the neighbouring countries. — Vasant
Press, Dehra Dun, 1007 pp. (Reprinted 1961).
Brunetti, E., 1923. Second revision of the Oriental
Stratiomyidae. — Records of the Indian Museum 25: 45-
180.
Brunetti, E., 1927: Notes on Malayan Diptera, with de-
scriptions of new species. — Journal of the Federal Malay
States Museums 13: 281-309.
Engel, E.O. & A. Cuthbertson, 1939. Systematic and bio-
logical notes on some brachycerous Diptera of southern
Rhodesia. — Journal of the Entomological Society of
South Africa 2: 181-195.
James, M.T., 1975. Family Stratiomyidae. In Delfinado,
M.D. & D.E. Hardy (eds.), A catalog of the Diptera of
the Oriental Region. Vol.II. The University Press of
Hawaii, Honolulu, 459 pp.
Lindner, E. 1928. Dr. L. Ziirchers Dipteren-Ausbeute aus
Paraguay: Stratiomyiiden. — Archiv fiir Naturgeschichte
(A) 22 (12): 94-103
McFadden, M. W., 1967. Soldier fly larvae in America
north of Mexico. — Proceedings of the United States
National Museum 121: 1-72.
Meijere, J. C. H. de, 1904. Neue und bekannte Siid-
Asiatische Dipteren. - Bijdragen tot de Dierkunde 17-18:
83-118.
Nagatomi, A., 1975. The Sarginae and Pachygasterinae of
Japan (Diptera: Stratiomyidae). — Transactions of the
Royal Entomological Society of London 126: 305-421.
Rozkosny, R., 1982/83. A biosystematic study of the
European Stratiomyidae (Dipteta). - Dr. W. Junk
Publishers, The Hague, Vol.1: 1-401, Vol. 2: 1-431.
Rozkosny, R. & D. Kovac, 1991. First description of the
male and the larva of Camptopteromyia fractipennis de
Meijere from Malaysia (Diptera: Stratiomyidae). —
Entomologica scandinavica 22: 297-304.
Wulp, F. M. van der, 1885. On exotic Diptera. — Notes
from the Leyden Museum 7: 1-15.
Received: 2 July 1993
Accepted: 9 February 1994
J. VAN TOL
National Museum of Natural History, Leiden, The Netherlands
iar ODONATA OE SULAWESTAND ADJACENT
ISLANDS
Part 3. The genus Macromia Rambur (Corduliidae)
Tol, J. van, 1994. The Odonata of Sulawesi and adjacent islands. Part 3. The genus Macromia
Rambur (Corduliidae). — Tijdschrift voor Entomologie 137: 87-94, figs. 1-25, table 1 [ISSN
0040-7496]. Published 15 July 1994.
The Macromia species of Sulawesi (Celebes) are detailed, and a key to the species is provided.
One new species, Macromia celebica sp. n., and the hitherto unknown female of M. irina
Lieftinck are described. The occurrence on Sulawesi of M. moore: fumata Krüger is considered
doubtful. The phylogenetic relationships of the Indo-Australian species of Macromia are ana-
lysed. The Sulawesi species are shown to have a sister group relationship to the monophyletic
group of the Papuan species. M. chalciope from Halmahera is considered the most recently
evolved taxon. These results are compared with recent reconstructions of the geological history
of the region.
J. van Tol, National Museum of Natural History, P. O. Box 9517, 2300 RA Leiden, The
Netherlands.
Key words. — Odonata; Corduliidae; Macromia; new species; phylogeny; biogeography;
Indonesia; Sulawesi; New Guinea.
' Part 2: Zoologische Mededelingen [1987] 61 (13): 160-176.
Hitherto, only two species of the rheophilic drag-
onfly genus Macromia Rambur have been mentioned
from Sulawesi (Celebes), viz. M. moorei fumata
Krüger and M. irina Lieftinck (Lieftinck 1950: 714-
716). Specimens of Macromia that were recently col-
lected in the southern part of Central Sulawesi, ap-
pear to belong to a species as yet undescribed. The
number of species reliably recorded from this island,
however, remains two, since the only record of
Macromia moorei fumata must be regarded doubtful
(see below).
The dragonflies of the virtually cosmopolitan ge-
nus Macromia are remarkable insects in many re-
spects. With more than 120 species, several of which
polytypic, Macromia is one of the largest genera of the
Anisoptera. They are especially speciose in the tropi-
cal areas of the Old World. In Europe (one species),
North America (nine species) and Australia (two spe-
cies), they are far less numerous. All other species are
confined to Africa and Southeast Asia.
Our present knowledge of the Macromia species of
Southeast Asia is mainly based on the studies of
Kriiger (1899) and especially Lieftinck (1929, 1950,
197 1a; see also 1954).
Due to their elusive behaviour, most species are
poorly represented in collections. In fact, many are
only known from the type series, often consisting of
one specimen only. Most species are fierce flyers. The
larvae seem to be quite stenotopic, so that larvae as
well as adults can only be found in very restricted
parts of streams. Notwithstanding their strong flying
capabilities, many species have small ranges, appar-
ently due to the fact that they seldom fly far from
their breeding sites.
In the present paper I describe the new species
from Sulawesi, provide some new observations on M.
irina, and give a key to species recorded from
Sulawesi. Finally, after having analysed the affinities
of the Sulawesi species to those of the Oriental and
the Papuan regions, I propose a hypothesis for a sce-
nario of the biogeographical history of this genus in
the Indo-Australian region.
SYSTEMATIC PART
Key to the Macromia species of Sulawesi
1. Outer margin of superior appendage of male with
well-developed tooth-like tubercle approximately
halfway (cf. fig. 18); vertex bilobed (cf. figs. 22,
23); thorax green with metallic lustre .................
D En OY A PRE LN M. moorei fumata
[Occurrence on Sulawesi doubtful].
— Outer margin of superior appendage of male
87
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 1-7. Macromia celebica sp. n. — 1, Head in frontal view; 2, Anal appendages of male, dorsal view; 3, Idem, left lateral
view; 4, Base of right hind wing, male; 5, Pterostigma of right fore wing; 6, Accessory genitalia of male, left lateral view; 7,
Fourth segment of penis, left lateral view.
somewhat curved inwards, in some specimens
with very small teeth along the margin, but espe-
cially in dorsal view not a conspicuous extero-lat-
eral tooth (figs. 2, 9); vertex dorsally smoothly
rounded, without a paired tubercle (figs. 1, 8);
thosaxqereenionbrownbe se ee eee 2
2. Appendix inferior distinctly shorter than superi-
ors (figs. 2, 3); coloration at least on synthorax
green with greenish blue metallic shine; frontal
side of head brownish black .... M. celebica sp. n.
— Appendix inferior approximately as long as
superiors (figs. 9, 10); coloration without any in-
dication of metallic lustre, but body generally
orange-brown; head orange-brown with some
middle-brown parts, but without conspicuous
brownishiblackstronsiseess ne M. irina
Macromia celebica sp. n.
(figs. 1-7)
Type material. — Holotype d. Indonesia. Southern
Central Sulawesi. Puncak Palopo. 600-1000 m.
88
October 1993. Leg. Yohan R. (JvT 6007) (MBBJ). —
Paratypes: 2d, same data (Jv I 5930-5931) (RMNH).
Description. — A medium-sized Macromia, with
green thorax and dark abdomen.
Male. Head with labium, labrum and clypeus
orange-brown, especially clypeus without yellow
markings, labrum with outer corners darker.
Pyramidal processes of frons high and subacute (fig.
1); frontal side very flat, more or less triangular, with
very fine transverse striae from upper-inner to lower-
outer corner of each side; frontal and upper parts of
frons shiny black with blue metallic lustre; frons
against eyes orange-brown as lower part of face; vertex
black with fine punctation, the paired dorsal tubercles
hardly discernable, the lateral angles tapering into
small projecting cones on top of which the lateral oc-
elli are situated (shape as in irina; westwoodii has
somewhat more pronounced dorsal tubercles), cov-
ered with thin setae. Occiput shiny black with very
fine punctation. Antennae brownish black.
Thorax. Brilliant metallic green, densely covered
with long and slender creamish white setae; antehu-
meral stripe yellow and distinct, somewhat tapering
dorsally, extending dorsally to ca. three-seventh of
mesepisternum, ventrally continuing over mesinfrae-
pisternum; yellow stripe over spiracle rather narrow,
tapering in width dorsally and nearly reaching the
upper margin of episternum. Pale stripe on latero-
ventral part of metepimeron covering anterior anteri-
or half (stripe absent in chalciope). Legs long and slen-
der, hind femur 10 mm, reaching to base of posterior
hamulus, colour brown, keel on fore leg very short,
ca. | mm, not reaching the apex, middle tibia without
keel, hind tibia with long keel from ca. 2 mm from
base of tibia to less than 1 mm from its apex.
Wings relatively short, fore wing with 17-18 Ax, 8
Px; hind wing with 11 Ax, 11-13 Px; fore wing with
7 Cux, hind wing with 5 Cux (incl. subtriangle); anal
loop 7-9 cells without central cell; anal triangle rela-
tively short (fig. 4; cf. M. chalciope, fig. 19); discoidal
triangle of hind wing very narrow; pterostigma (fig.
5) reddish brown, anterior side ca. 2.0 mm.
VAN TOL: Macromia of Sulawesi
Abdomen brownish black with some blue-green
metallic lustre especially on first few segments; yellow
spot on anterior side of segment 7; general shape an-
teriorly slender, especially segment 7-9 significantly
widened; tergites ventro-anteriorly somewhat paler,
dark or middle-brown. Spot on segment 7 consisting
of a complete ring covering anterior one-fourth of
segment, medio-dorsally extending posteriorly for an-
other one-eighth segment length and ca. one-third
the segment’s width. Segment 10 rather short, mid-
dorsal process not spine-like, but consisting of a low
triangular-shaped subterminal tubercle. Appendages
(figs. 2-3) with superiors in dorsal view rather slender,
significantly longer than inferior, basal two-fifths
rather straight, then innerside slightly tapering poste-
riorly and outerside strongly tapering, caudalmost
part somewhat curved outwards; no trace of a lateral
tubercle on superior appendage.
Secondary genitalia: anterior side margin of tergite
2 with dense setae, a mixture of strong and soft setae,
longer than in zrina; also tuft of setae on ventral tu-
Figs. 8-15. Macromia irina Lieftinck. — 8, Head in frontal view; 9, Anal appendages of male, dorsal view; 10, Idem, left later-
al view; 11, Base of right hind wing, male; 12, Pterostigma of right fore wing; 13, Accessory genitalia of male, left lateral view;
14, Fourth segment of penis, left lateral view; 15, Valvula vulvae, female.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
bercle of tergite longer; genital lobe directed posteri-
orly, elongate, and tapering towards the top, but tip
not acute; posterior hamulus glossy brown, basally
coloured brown, the straight outer part pale brown,
and the endhook reddish brown, curved strongly (90°)
inwards, endhooks in ventral view slightly diverging,
the tip reaching ca. two-thirds of genital lobe.
Measurements. — Abdomen, incl. appendages 48
(paratypes 46-48) mm, hind wing 42 (paratypes 42-
43) mm
Female. — Unknown.
Differential diagnosis. — Differs clearly from the
only other Sulawesi Macromia by the presence of
green metallic lustre, the black markings on the frons,
the presence of an antehumeral stripe; structurally the
short inferior appendage, and shape of abdominal
segment 10 are diagnostic. Both Sulawesi species can
be distinguished from other Indo-Australian
Macromia by the intermediate sized pterostigma, the
very narrow discoidal triangle of the hind wing, and
especially the rounded (not bilobed) vertex. Also, the
superior appendages lack an extero-lateral tooth, but
are not particularly slender. The Papuan Macromia
have extremely slender superiors with or without an
extero-lateral tooth (fig. 20), while the Oriental spe-
cies have rather solid appendages with a more or less
conspicuous extero-lateral tooth (fig. 18).
Remarks. — Although this species differs in many
respects from M. irina, I consider it the sister species
of M. irina. This view is based on shared, presumably
apomorphous, character states in the structure of ver-
tex and superior appendage of male (see below).
Distribution. — Only the southwestern part of
Central Sulawesi. Collected between 600-900 m.
Macromia irina Lieftinck
(figs. 8-15)
Macromia irina Lieftinck, 1950: 714-716. Holotype male ‘S
Celebes, foot of Mt Lompobatang, Borong Rapao, 800
m, 12.viii. 1949, leg. A. Diakonoff in RMNH [examined].
— Lieftinck 1971b: 94 [note on type]; Askew et al. 1989:
118 [records Northern Sulawesi, Dumoga Bone National
Park].
Diagnosis. — Unmistakable among Indo-Australian
Macromia: head, thorax and abdomen brown without
any metallic lustre. Structural characters: wing base
with anal angle sharp, but not angulated (fig. 11); the
anal triangle rather short (long in moore fumata),
membranula ending circa halfway between anal cross-
vein and base of anal triangle; pterostigma rather
short (fig. 12), circa twice as long as high (more than
three times in moorei fumata); anal appendages of
male with superiors with rather smooth outer margin,
without any indication of tooth (figs. 9-10) (cf. moo-
90
rei fumata and westwoodii); inferior appendage ap-
proximately as long as superiors (clearly shorter in ce-
lebica).
Description. — Little can be added to the extensive
description of the male in Lieftinck (1950). Live
colour of eye dark green. Since Lieftinck did not pro-
vide any illustration, I have added here drawings of
the head, wing base, anal appendages and secondary
genitalia (figs. 8-14). The up to now undescribed fe-
male was collected in Southeast Sulawesi.
Female. — General coloration as the male, and
structurally very similar to the male, although more
robust. Head and thorax not differing from male.
Wings somewhat broader, especially at base; mem-
branula fully opaque white or only greyish in most
narrow part; rusty brown base of wings more exten-
sive than in male, in costal and subcostal area of fore
and hind wing reaching to first Ax or even somewhat
beyond; some less extensive rusty brown at base of
medio-anal space; Arculus at Ax2 in fore wing and
just beyond Ax2 in hind wing; fore wing 16-17 Ax, 9
Px 7 Cux; hinde wing 10212 Ax NDR Cuxe
Abdomen of the coloration (‘Mikado-brown’) as
male; segment | dorsum dark brown and lateral parts
brown; segment 2 brown, somewhat darker above
than laterally, as the male with two narrow transverse
yellow streaks, one over transverse carina halfway ab-
dominal segment and the other pair more oblique
running just before posterior margin; segment 3 ante-
riorly brown to transverse carina, somewhat darker
above, posteriorly dark brown except for space just
before segment 4; segments 4-7 similar to segment 3,
but dark patch anterior to transverse carina gradually
increasing in size, and posterior paler brown ring also
increasing from segment 4 to 7; segment 8 brown
with dorsal carina black and some darkening against
dorsal carina; segment brown, dorsal carina dark;
segment 10 brown; appendages middle brown, rather
short, acute. Valvula vulvae short, approximately four
times as wide as long, with wide triangular emargina-
tion, the tips just bifid, somewhat curved inwards
(fig. 15).
Other material examined. — Sulawesi Tenggara. S
of Sanggona: Gunung Watuwila. Sunggai Lalondu-
wasi near Centipede camp. c. 1050 m. 2-4 Nov 1989
(J. van Tol) 36 19 (JvT 5932-5935); Sulawesi
Tengah, ca. 10 km WNW Palopo near Tojambu.
800-1000 m, c. 2°56’S 120°07’E, Jul/Aug 1991 (Yo-
han R.), 12 (JvT 5936), all in RMNH. I have also
studied one of the specimens collected by Askew et al.
(1989): 1 6 , in RSME.
Remarks. — I collected this species myself along a
small, shaded stream in a steep valley in the
Mengkoko mountain area. The insects appeared im-
mediately after sunrise, and were active for approxi-
VAN TOL: Macromia of Sulawesi
Figs. 16-23, Macromia species. 16-18, 22, M. westwoodii, 19-21, 23, M. chalciope. — 16, 19, Base of right hind wing, male;
27, 21, Pterostigma of right fore wing; 18, 20, Anal appendages of male, dorsal view; 22-23, Vertex in frontal view (fig. 23
dorsal-most part only).
mately one hour. Males as well as females flew low
over the water up and down the stream, more or less
in a straight line. With a rather slow and small wing
beat, they were not extremely fast. The specimens
could be collected at a site where a big log of more
than one meter diameter was laying in the stream.
Macromia specimens crossed the log at just a few cen-
timeters height to return immediately to the water
surface. The female was caught on a dark day at 16.30
h, i.e. 1.5 hours before sunset.
Macromia moorei fumata Krüger
Macromia fumata Kriiger, 1899: 333-335 (Java) [not exami-
ned].
Macromia moorei fumata. — Lieftinck 1950: 678 (key, men-
tioned Celebes); Lieftinck 1954: 119, 185 (references,
geographical distribution); Lieftinck 1971a: 29 (geo-
graphical distribution).
Remarks. — I have been able to check the specimen
(in SMFD) at which Lieftinck’s notice of this taxon on
Sulawesi was based. Although this specimen can be
assigned to M. moorei fumata without doubt, | con-
sider its occurrence on Sulawesi still doubtful. The
‘Lompo Battau’ labels of Fruhstorfer have on many
occasions proved to be incorrect (see van Tol 1987:
163). M. moorei fumata is rather common on Java,
where Fruhstorfer also collected during his long jour-
ney through the former Dutch Indies.
Material examined. — Lompa Battau, S. Celebes,
III. 1896 (H. Fruhstorfer) ‘16764 (in green ink) 14
[label in Ris’ hand] [identified by M.A. Lieftinck as
Macromia moorei fumata, Jan. 1928].
PHYLOGENETIC RELATIONSHIPS
Introduction
The so-called Papuasian représentatives of Macro-
mia share at least four characters (Lieftinck 1952,
1971a), viz., an exceptionally small discoidal triangle
of the hind wing, a minute pterostigma of ca. one
mm, a sharply acute anal angle of the hind wing in
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Table 1 Character matrix for taxa of Macromia of the Indo-Australian archipelago.
(1) extero-lateral tooth superior appendage
(2) vertex
(3) hamulus
(4) superior appendages
(5) pterostigma
(6) postclypeus
(7) antehumeral stripe
(8) anal angle
(9) discoidal cell
(10) general coloration
(11) dorsal spine abdominal segment 10
(12) curvature superior appendages
Character 1 2 3 4 5
Species
amymone 1 1 2 2 3
celebica DD 2 1 1 2
chalciope 3 1 2 2 3
trina 2 2 1 1 2
melpomene 1 1 2 Dy, 3
moorei fumata 1 1 1 1 1
terpsichore 3 1 2 2 3
westwoodii 1 2 1 1 1
the male, and long and slender genital hamules in the
male. Lieftinck (1971a: 30-32) distinguishes three
groups among the Papuan Macromia species, viz. M.
terpsichore, M. melpomene, and M. chalciope, with the
last one consisting of one species only.
The species assigned to these groups, which the ex-
ception of viridescens described from Cape York
(Australia), are confined to the islands of New
Guinea (astarte Lieftinck, celaeno Lieftinck, eurynome
Lieftinck, melpomene Ris, terpsichore Foerster, virides-
cens Tillyard (= amymone Lieftinck)), the Bismarck
Archipelago (lachesis Lieftinck), Waigeu (euphrosyne
Lieftinck, sophrosyne Lieftinck) and Misool (hermione
Lieftinck). M. chalciope Lieftinck is restricted to
Schouten Is, and the Moluccan islands of Halmahera
and Bacan (Lieftinck 1971a). No species of
Macromia have been mentioned from any of the oth-
er Moluccan islands up to now. According to
Lieftinck (1952: 460) M. chalciope can be considered
‘an immediate derivative of the eastern cluster’, i.e.
that the sister-species of chalciope could be discovered
among the species of New Guinea and adjacent is-
lands.
The Malay peninsula and the Greater Sunda
Islands are inhabited by 15 species of Macromia (see
Lieftinck 1971a: 29). Although Lieftinck (1929) at-
tempted to distinguish species groups in the western
Malesian species, this author (1950: 716) had to ad-
mit that the groups had an artificial character.
Variation in morphology among the species of west-
92
12 (present/absent)
12 (bilobed/rounded)
12 (shorter/longer lobus)
12 (normal/very slender)
123 (long/intermediate/ very short)
12 (unicolorous/yellow markings)
12 (present/absent)
12 (sharp/angulate)
12 (wide/narrow)
12 (green/brown)
123 (slender/absent/broad)
123 (curved inward/outward/straight)
6 7. 8 9 10 11 12
2) 2 2 2) 1 2 2
1 1 1 2 1 2 À
1 1 22 2 1 1 3
1 2 1 2 2 2) 2
1 2 2 2. 1 1 3
1 2 1 1 1 ?) 1
2 1 2 2 1 2) 3
1 1 1 1 1 3 2
ern Malesia is more prominent than that among the
Papuan representatives.
Finally, the Philippine islands are inhabited by at
least one endemic species, viz. M. negrito Needham &
Gyger from Luzon. The widespread M. cincta is re-
ported from Palawan (Lieftinck 1929: 92), and Tawi
Tawi (Lieftinck 1974), while also M. gerstaeckeri may
amymone
chalciope
terpsichore
melpomene
celebica
trina
westwoodii
moorei fumata
Fig. 24. Tree for Macromia species of the Indo-Australian
archipelago.
VAN TOL: Macromia of Sulawesi
Fig. 25. Phylogenetic relationships of Macromia species (groups) represented on the map of Malesia.
occur in the Philippines (Lieftinck 197 1a: 29).
I present here the results of a preliminary investiga-
tion of the phylogeny of the Macromia of the Indo-
Australian region, mainly based on the characters
used by Lieftinck to establish his Papuan species
groups, or to distinguish the Oriental and Papuan
groups. As outgroups I have added M. moorei fumata
Krüger and M. westwoodii Selys, both considered
closely allied to M. irina (Lieftinck 1950). For the
present study only a limited number of the Papuan
species have been included. Unfortunately, M. negri-
to Needham & Gyger from Luzon, was not available
for study. A more extensive phylogenetic study, in-
volving more of the Oriental as well as the Papuan
species, has to wait for the future.
Methods and results
The matrix (table 1) was analysed with PAUP, ver-
sion 3.1.1 (Swofford 1993), using “exhaustive search’.
Macromia moorei fumata was designated as outgroup,
but the results appeared to be similar when M. west-
woodii was selected as such. The analysis resulted in
one tree of 20 steps (CI=0.800, RI=0.818).
This tree (fig. 24) shows that all Papuan species
form a monophyletic group, with the Sulawesi species
as sister group. The Papuan groups as defined by
Lieftinck (197 1a) are not corroborated by the present
analysis. It is of interest to notice the sistergroup rela-
tionship of M. terpsichore from NE New Guinea, and
M. chalciope from Halmahera.
Homoplasies in the present tree expecially occur in
two non-structural characters, viz. character 7 (ante-
humeral stripe) and to a lesser extent character 6
(postclypeus).
Discussion and biogeography
The results of the phylogenetic analysis partly con-
firm earlier suggestions by Lieftinck on relationships
among Indo-Australian Macromia species. Lieftinck
(1950) considered M. irina from Sulawesi closely re-
lated to M. moorei and M. westwoodii, both rather wi-
despread in Southeast Asia, including Borneo. The
rather solid build of the superior appendage, in com-
bination with the absence of an extero-lateral tooth,
ranks the Sulawesi representatives in an intermediate
position between the Oriental and the Papuan spe-
cies. This intermediate position can also be recogni-
zed in the shape of the pterostigma.
The rather distant relationships of the Sulawesi and
Halmahera species are of interest in a biogeographical
context. Although the geological history of Wallacea
and Halmahera is still under debate (e.g. Daly et al.
1991, Honza 1991), there is general agreement that
there is no shared history for the North Moluccas
(Halmahera) and Sulawesi. Reconstructions prior to
10-15 million years ago (Ma) are highly uncertain be-
cause of the very complicated plate movements. In
the reconstruction of Daly et al. (1991) (see also
Nichols & Hall 1991) Halmahera was formed at the
margin of the Pacific and the Philippine plates, north
of New Guinea, with a westward progression from c.
5 Ma to its present position. At least some parts of
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Sulawesi have a much longer (65 million years) histo-
ry above sea level on a far more western position. In
other words, from approximately 60 to 5 Ma, the
Oriental region + Sulawesi have shared (in some re-
spects) a common history, while Halmahera shares its
history mainly with New Guinea. The close contact
of the Oriental and Papuan region is more recent
than 5 Ma.
Given these phylogenetic relationships, and our
present knowledge of the geological history of the
area, the following historical scenario may be outli-
ned. All Macromia species in the Indo-Australian re-
gion originate from an Oriental ancestor. The ance-
stor of the Sulawesi + Papuan species groups possibly
inhabited Sulawesi, while the ancestor of the
Halmahera M. chalciope + New Guinea M. terpsicho-
re species groups apparently lived in north(eastern)
New Guinea. The occurrence of two relatively remo-
tely allied species on Sulawesi and Halmahera can be
attributed to the westward movement of Halmahera
during the last ten million years.
ACKNOWLEDGEMENTS
Field work in Sulawesi was supported by the
Zoological Museum at Bogor. The kind help of the
director of the museum, drs. Mohammad Amir, and
his staff, as well as the staff of Lembaga Ilmu Pengata-
huan Indonesia (Indonesian Academy of Sciences) is
gratefully acknowledged. The expert knowledge of
Sulawesi of Mr. Baharudin (Bantimurung) was of
great help for successful collecting. Dr. H. Schròder
kindly arranged the loan of material in SFMD. I also
thank Dr. K. Wilson (Hongkong) for his comments
on a draft of this paper.
REFERENCES
Askew, R.R., G.G. Cleland, D.A.L. Davies & T.W.
Harman, 1989. A report on a collection of Odonata from
North Sulawesi, Indonesia. — Tijdschrift voor Entomolo-
gie 132: 115-121, fig. 1, tables 1-2.
Daly, M.C., M.A. Cooper, I. Wilson, D.G. Smith &
B.G.D. Hooper, 1991. Cenozoic plate tectonics and ba-
94
sin evolution in Indonesia. — Marine and Petroleum
Geology 8: 2-21, figs. 1-15.
Honza, E., 1991. The Tertiary arc chain in the Western
Pacific. — Tectonophysics 187: 285-303, figs. 1-5.
Kriiger, L., 1899. Die Odonaten von Sumatra. III. Theil. a.
Familie Libelluliden (with supplement: Neue Cordulinen
von Java und Borneo. — Stettiner Entomologische Zei-
tung 60: 321-338.
Lieftinck, M. A., 1929. A revision of the known malaysian
dragonflies of the genus Macromia Rambur, with com-
parative notes on species from neighbouring countries
and descriptions of new species. — Tijdschrift voor
Entomologie 72: 59-108, figs. 1-22.
Lieftinck, M. A., 1950. Further studies on Southeast Asiatic
species of Macromia Rambur, with notes on their ecolo-
gy, habits and life history, and with descriptions of larvae
and two new species (Odon., Epophthalmiinae). —
Treubia 20: 657-716.
Lieftinck, M. A., 1952. On the Papuasian representatives of
the genus Macromia Ramburm with descriptions of five
new species and some larval forms (Odon.). — Treubia 21:
437-468, p. 1-22.
Lieftinck, M. A., 1954. Handlist of Malaysian Odonata. —
Treubia 22 (Supplement): i-xiii + 1-202, map.
Lieftinck, M. A., 1971a. Studies in Oriental Corduliidae
(Odonata), I. — Tijdschrift voor Entomologie 114 (1): 1-
63, figs. 1-50.
Lieftinck, M. A., 1971b. A catalogue of the type-specimens
of Odonata preserved in The Netherlands, with a supple-
mentary list of the Odonata types described by Dutch sci-
entists deposited in foreign institutional collections. —
Tijdschrift voor Entomologie 114 (2): 65-139 + plate 1-
Ve 9
Lieftinck, M. A., 1974. Dragonflies collected by the Noona
Dan Expedition in the southwestern Philippine Islands
(Insecta, Odonata). — Steenstrupia 3: 111-147, figs. 1-26.
Nichols, G. J. & R. Hall, 1991. Basin formation and
Neogene sedimentation in a backarc setting, Halmahera,
eastern Indonesia. — Marin and Petroleum Geology 8 (1):
50-61, figs. 1-9.
Swofford, D. L., 1993. paup: Phylogenetic Analysis Using
Parsimony. Version 3.1.1. — Illinois Natural History
Survey, Champaign.
Tol, J. van, 1987. The Odonata of Sulawesi and adjacent is-
lands. Part 2. The genus Diplacina Brauer on Sulawesi. —
Zoëlogische Mededelingen 61: 160-176, figs. 14-39.
Received: 2 February 1994
Accepted: 14 April 1994
JOHN S. WEAVER III’ & HANS MALICKY’
! University of New Hampshire, Durham, U.S.A. * Biologische Station Lunz der Österreichischen
Akademie der Wissenschaften, Lunz am See, Austria.
THE GENUS DIPSEUDOPSIS WALKER FROM ASIA
GUICGHORTERA: DIPSEUDOPSIDAE)
Weaver, J. S., III & H. Malicky, 1994. The genus Dipseudopsis Walker from Asia (Trichoptera:
Dipseudopsidae). — Tijdschrift voor Entomologie 137: 95-142, figs. 1-55. [ISSN 0040-7496].
Published 15 July 1994.
The genus Dipseudopsis Walker from Asia is revised, and a phylogeny of Hydropsychoidea is
provided showing the relationships of its family taxa and the genera of Dipseudopsidae.
Presently 36 Asian species of Dipseudopsis are recognized, having a distribution including
Pakistan, India, Sri Lanka, Nepal, Bangladesh, Burma, Thailand, Cambodia, Vietnam, China,
Japan, Philippines, Malaysia and Indonesia, including Sumatra, Java, Borneo, and Sulawesi. Six
new species are described, Dipseudopsis adiaturix sp. n., D. flinti sp. n., D. lucasi sp. n., D. mal-
aïsei sp. n., D. martynovi sp. n., and D. schmidi sp. n., and one new subspecies, D. robustior an-
damanensis ssp. n. The identities of several problematic species are resolved, with 16 species
names recognized as new junior synonyms (cf. species checklist p.102). D. nervosella Ulmer
stat. n. is elevated from former subspecies status. New descriptions are provided for all species,
including figures of the male hind mesoapical tibial spur (the modified spur), the male genital-
ia, and other notable characteristics of the head, thorax and wings, except two species recog-
nized as nomina dubia, D. onychophora Navas, and D. orientalis (Navas). New lectotypes are
designated for D. contorta Banks, D. morosa Banks and D. triclavata Martynov. A pictorial at-
las is provided for rapid species identification. The genus Eodipseudopsis Marlier is removed
from the Dipseudopsidae.
Correspondence: J. S. Weaver III, Department of Entomology, University of New Hampshire,
Durham, NH, 03824, U. S. A.
Key words. — Asia, Dipseudopsis, Dipseudopsidae, Hydropsychoidea, phylogeny, systematics,
Trichoptera.
The genus Dipseudopsis Walker exhibits a number
of characteristics that are quite remarkable among the
Trichoptera. The adults are the only members of the
suborder Annulipalpia Martynov having siphoning
mouthparts, with the lacinia forming a proboscis, a
parallel modification of the galea in the higher
Lepidoptera. Sexual dimorphism is conspicuous in
the genus, with the males having highly contrasting
colour patterns in their wings and bodies, and the fe-
males less striking in comparison. Also, males have
the mesoapical spur of the hind tibia modified, often
twisted and/or branched. The function of the modi-
fied spur is unknown, but it has been used almost ex-
clusively for species determinations, to the extent that
customary descriptions of the male genitalia often
have been omitted from species descriptions. The
convention of using the modified spur as the basis for
species descriptions was established by Ulmer
(1907b: 180): “Bei Dipseudopsis sind die Genitalien
aller Species ganz gleich’ (i.e: In Dipseudopsis the gen-
italia of all species are totally alike). Hence, the mod-
ified spur became the primary characteristic used to
recognize species; e.g. this is the main distinguishing
character in every couplet of Ulmer’s (1951) key to
eleven species of Dipseudopsis of the Sunda Islands.
The precise orientation of the modified spur must
be made apparent for species identification, because
slight rotations of the tibia will show different per-
spectives of the spur. Some of the previous species de-
scriptions based on the modified spur were ambig-
uous, because the figure of the spur did not show the
remainder of the leg, and hence the actual orientation
of the spur was uncertain. This problem accounts for
several of the synonyms that we discovered in the
course of this study. The figures of the spur herein are
usually of the left leg and include the adjacent apic-
olateral spur and a portion of the tibia and tarsus to
provide proper comparison of other specimens in the
same perspective.
It seems somewhat ironic that the taxonomy of this
genus, which exhibits so many fascinating character-
95
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Table 1. List of character states. 0 = Plesiomorphic in respect to the outgroup Philopotamidae. 1 = Synapomorphic character.
1. Adult dorsal ocelli present = 0; ocelli absent = 1 (cf. Schmid 1980).
ho
Schmid 1980).
. d segment IX completely sclerotized cylinder = 0; IX jawlike with articulation between tergum & sternum = 1 (cf.
2 segment X without lateral papillae = 0; X with lateral papillae = 1 (cf. Schmid 1980).
2 segment IX normal = 0; IX long completely sclerotized cylinder = 1 (cf. Schmid 1980).
. Larva spinneret short = 0; spinneret longer than other mouthparts = 1 (cf. Wiggins 1977).
. à inferior appendages 2-segmented = 0; 1-segmented = 1 (cf. Schmid 1980).
. Adult bases of antennae distant = 0; bases close together = 1 (cf. Crichton 1957).
3
4
5
6. 2 sternum VIII normal, undivided = 0; sternum VIII divided into a pair of lateral lobes = 1 (cf. Schmid 1980).
7
8
9
. Larva tarsi nearly cylindrical, pretarsal claws long = 0; tarsi flattened, claws short = 1 (Ulmer 1957, Gibbs 1968, Wiggins
1977, Wells & Cartwright 1993).
10. Larval retreat differs from the following = 0; larva with branched tube-dwelling and filter-feeding behaviour as in
Phylocentropus (Wallace et al. 1976, Wiggins 1977).
11. Adult labial palps present = 0; absent = 1 (Ulmer 1951).
12. Adult pronotum normal = 0; pronotum enlarged and with deep median fissure = 1.
13. Adult with head and thorax setous = 0; head and thorax glabrous = 1.
14.. Adult tegula knoblike = 0; tegula fattened like epaulets = 1.
15. Adult with galea short, vestigial = 0; modified into proboscis = 1.
16. Larva labrum expanded and more membranous = 1, autapomorphy for Philopotamidae (cf. Wiggins 1977).
17. Larva trochantin broad, either with blunt apical angle or truncate = 1, autapomorphy for Psychomyiidae (cf. Wiggins
1977).
18. Larva with mesopleural lobes = 1, autapomorphy for Xiphocentronidae (cf. Wiggins 1977).
19. Larva with tibiae and tarsi of legs fused = 1, autapomorphy for Xiphocentronidae (cf. Wiggins 1977).
20. Larva with ventral thoracic gills whorled = 1, autapomorphy for Hydropsychidae (cf. Wiggins 1977).
21. Larva epidermis with modified setae, especially on sternum VII] & IX and tergum = 1, autapomorphy for
Hydropsychidae (cf. Wiggins 1977).
22. Larva head with ventrolateral bands of transverse ridges used in stridulation = 1, autapomorphy for Hydropsychidae (cf.
Wiggins 1977).
23. Larval head and body flattened dorsoventrally = 1, autapomorphy for Ecnomidae (cf. Lepneva 1964).
24. Larva lateral fringe dense = 1, autapomorphy for Ecnomidae (cf. Lepneva 1964).
istics, should have become so confused at the species
level. This is chiefly because Banks, Martynov, Navas,
and Ulmer from 1905 to 1937 proposed descriptions
of 25 different species from Asia and the species de-
scriptions by Banks and Navas are poor. Another fac-
tor might be that specimens of Dipseudopsis are not
particularly abundant in museum collections, even
though adults are attracted to light traps.
The primary objective of this work is to clarify the
taxonomy of the Asian species of Dipseudopsis. To ac-
complish this we have examined the types of most of
the species as well as other specimens from several
museums and private collections. The species descrip-
tions have been expanded beyond the characteristics
of the modified spur to include colouration of the
body and wings and illustrations of the male genitalia.
Hence, most of the redescriptions of species herein
provide the first illustrations of the male genitalia. We
recognize 36 species of Dipseudopsis from Asia, 30
previously described species and six new species, and
one new subspecies. We also recognize the names of
16 species and one subspecies as new junior syno-
nyms. Two additional Asian species, D. onychophora
Navás and D. orientalis (Navás), are recognized as no-
mina dubia, because both were poorly described and
their types could not be located for examination.
96
DEPOSITORIES
The depositories of material are abbreviated as fol-
lows: BMNH, Natural History Museum (formerly the
British Museum of Natural History), London; BPBM,
Bernice Pauahi Bishop Museum, Honolulu; CASC,
California Academy of Sciences, San Francisco; CNCI,
Canadian National Collection, Ottawa; CLDD, collec-
tion of D. G. Denning (now deposited at CASC);
CLHM, collection of H. Malicky; CLJW, collection of J.
S. Weaver III; MCZC, Museum of Comparative
Zoology, Harvard University, Cambridge; MNHN,
Muséum National d’Histoire Naturelle, Paris; MZBS,
Museu de Zoologia, Barcelona; MZLU, Museum for
Zoology, University of Lund; NHMw, Naturhistori-
sches Museum Wien; NHRS, Naturhistoriska Riksmu-
seet, Stockholm; NZSI, National Zoological Collec-
tion, Zoological Survey of India, Calcutta; RMNH,
Nationaal Natuurhistorisch Museum (formerly Rijks-
museum van Natuurlijke Historie), Leiden; SOFM,
National Museum of Natural History, Sofia; UOPJ,
Entomological Laboratory, University of Osaka
Prefecture, Sakai; USNM, Smithsonian Institution,
United States National Museum of Natural History,
Washington; ZRAS, Zoological Institute of the Rus-
sian Academy of Sciences, St. Petersburg; ZMAN, Zoo-
logisch Museum, Amsterdam; ZMHB, Zoologisches
Museum der Humboldt-Universität Berlin; ZMUC,
Zoologisk Museum Kobenhavn; ZMUH, Zoologisches
Museum der Universität Hamburg; and ZSMC,
Zoologische Staatssammlung München.
METHODS
Taxonomy
The morphological terminology of wing venation
and genitalic characters mostly follows that of Schmid
(1980), and Schuster & Hamilton (1984). The species
synonymies are complete listing all original species de-
scriptions, but subsequent references listed in Fischer’s
Trichopterorum Catalogus (1962, 1972) are not inclu-
ded. Locality names in current use are provided for spe-
cimens examined, where old names taken from labels
are in quotes. For the type data, the term ‘holotype’ is
used when the type specimen has actually been labelled
or cited as such, otherwise it is listed as ‘type’. An aste-
risk (*) designates types of previously described species
examined by the authors.
Phylogenetics
Hennig86 (Farris 1988) is used for phylogenetic
analysis to determine the relationship of
Dipseudopsidae within the Hydropsychoidea s. str.
Weaver, and the relationships of the proposed di-
pseudopsid genera. Twenty four character states (ta-
ble 1) are selected, and with polarity determined for
twelve taxa, using Philopotamidae Stephens as an
outgroup, the data matrix (table 2) is produced.
These data are analyzed using the ‘mhennig*’ com-
mand of Hennig86 for calculating trees. All charac-
ters have default values, weight = 1, active and additi-
ve, except weight = 2 for character 6.
WEAVER & MALICKY: Dipseudopsis from Asia
DIPSEUDOPSIDAE ULMER
The family Dipseudopsidae is included in the sub-
order Annulipalpia Martynov, in the infraorder
Curvipalpia Weaver (Weaver 1984, Weaver & Morse
1986). The family name, Dipseudopsinae Ulmer
(1904b), including only the nominative genus, was
first proposed as a subfamily of the Hydropsychidae
Curtis, but was later placed within the Polycentropo-
didae Ulmer (1906), as it has been recognized by
several others (Tsuda 1942, Marlier 1962, Ross &
Gibbs 1973, Wiggins 1977, Schuster & Hamilton
1984). However, some workers in the past have re-
cognized Polycentropodinae and the Dipseudopsinae
as subfamilies of Psychomyiidae Curtis (Ross 1956,
Ross & Kingsolver 1959), and more recently, some
still recognize the Polycentropodinae as such (Flint
1991). The Dipseudopsidae were first given full fami-
ly status by Ross (1967), and later concurred by
Gibbs (1968), Wiggins (1982), Weaver (1984), Scott
(1985), Weaver & Morse (1986), and Wells &
Cartwright (1993).
The genus Protodipseudopsis Ulmer (1909) from
the Afrotropical Region was the first addition to the
subfamily, a relationship later confirmed by Ross &
Kingsolver (1959). Two additional genera from the
Afrotropical Region were placed in the Dipseudo-
psinae, Eodipseudopsis Marlier (1959) by original des-
ignation, and Limnoecetis Marlier (1955) originally
placed in the Leptoceridae Leach, but later moved to
the Dipseudopsinae (Marlier 1961). Ross & Gibbs
(1973) transferred Phylocentropus Banks (1907) from
Polycentropodinae to Dipseudopsinae, based pre-
dominantly on synapomorphic larval characteristics.
They recognized only three genera in the Dipseudo-
psinae, Dipseudopsis, Phylocentropus, and Protodipseu-
dopsis, and did not consider the status of Eodipseu-
dopsis and Limnoecetis. However, Schmid (1980) later
Table 2. Data matrix for the families of Hydropsychoidea and genera of Dipseudopsidae, including outgroup Philopotamidae.
Character
Il
12345 67890
Taxon
Philopotamidae 00000 00000
Psychomyiidae 11011 00000
Xiphocentronidae 11011 00000
Hydropsychidae 11100 10000
Polycentropodidae 11100 11000
Ecnomidae 11100 11000
Dipseudopsis 11101 01111
Protodipseudopsis 11101 01111
Phylocentropus 11101 01111
Limnoecetis 1110? 0112?
Hyalopsyche 11101 01111
Hyalopsychella 1122? Dit 228
11111 11112 2222
12345 67890 1234
00000 10000 0000
00000 01000 0000
00000 00110 0000
00000 00001 1100
00000 00000 0000
00000 00000 0011
01111 00000 0000
01110 00000 0000
00000 00000 0000
01110 POLE HE
10000 00000 0000
10000 OO DIE GORE
2/7
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
made the novel assignment of placing Phylocentropus in
the family Hyalopsychidae Lestage, on the basis of
adult characteristics shared with the genus Hyalopsyche
Ulmer (1904b). Wells & Cartwright (1993) recently
concurred with the position of Ross and Gibbs, and
based on the morphology of the larva and female of
Hyalopsyche, added this genus to the Dipseudopsidae,
and accordingly proposed the suppression of the family
group name Hyalopsychidae. Our phylogenetic anal-
ysis tests the classifications of Ross & Gibbs and Wells
& Cartwright, versus that of Schmid.
Results
Phylogeny. — The analysis of the data matrix via
Hennig86 generated three trees with different topolo-
gies, having length 28, CI 0.89, RI 0.91 (fig. 55). All
three trees depict Dipseudopsidae as a monophyletic
group, comprising six genera. The first tree has the fe-
west number of ancestors, having 18 nodes, and is
also identical to the strict consensus tree (fig. 56) of
the three trees. The consensus tree depicts ((Psycho-
myiidae, Xiphocentronidae Ross) ((Hydropsychidae,
Ecnomidae Ulmer, Polycentropodidae) Dipseudo-
psidae)), thus supporting monophyly of the Hydro-
psychoidea. The clade comprising Hydropsychidae,
Polycentropodidae, Ecnomidae is recognized as the
sister group of the Dipseudopsidae. A few differences
are noted among the three original trees: 1) trees 0
and 2 depict the trichotomy (Hydropsychidae, Ecno-
midae, Polycentropodidae), which is resolved as (Hy-
dropsychidae (Ecnomidae, Polycentropodidae)) in
tree 1; 2) trees 0 and 1 each have a basal and a distal
trichotomy among the genera of Dipseudopsidae,
both of which are resolved in tree 2.
Characters. — The combination of characteristics of
the female genitalia as exhibited in the Dipseudo-
psidae (having synapomorphy 2, lateral papillae of
segment X present, but lacking synapomorphy 6, ab-
dominal sternite VIII not subdivided into lateral
lobes) does not represent an apomorphic state, but
only a unique combination of characteristics which
happens to define the family very well, but only in a
taxonomic sense. Synapomorphy 7, inferior appen-
dage one-segmented is a homoplasy in Polycentro-
podidae and Ecnomidae. Three synapomorphies 8-
10 support monophyly of Dipseudopsidae, and the
first of these, antennal bases close together, is not
unique, but within the Hydropsychoidea is also
present in Hydropsychinae and Macronematinae
(Hydropsychidae). However, since this characteristic
is absent for the most part in Arctopsychinae and
Diplectroninae, it is not part of the groundplan of
Hydropsychidae, and thus is a homoplasy. A more ac-
curate description of this condition as a homologue is
98
that the antennal scapes of Dipseudopsidae are not
merely close together but rather are so enlarged that
they occupy most of the space between the eyes along
the anterodorsal margin of the head (fig. 13).
Conclusions. — Monophyly of the Dipseudopsidae,
including Dipseudopsis, Hyalopsyche, Hyalopsychella
Ulmer (1930), Zimnoecetis, Phylocentropus, and
Protodipseudopsis, is unequivocal. Two clades are rec-
ognized among these genera, (Hyalopsyche + Hyalo-
psychella) supported by synapomorphy 11 (cf. table
1), and (Dipseudopsis + Protodipseudopsis + Limnoe-
cetis) supported by synapomorphies 12-14. Hence,
the sister group of the genus Dipseudopsis is unclear.
The adult of Zimnoecetis, a monotypic genus endem-
ic to Lake Tanganyika, are quite remarkable, having
unique adaptations to walk on water. The striking au-
tapomorphies of Limnoecetis present a formidable
challenge in resolving the trichotomy (Dipseudopsis +
Protodipseudopsis + Limnoecetis). One obvious predic-
tion based on our phylogeny, is that the unknown lar-
val forms of Hyalopsychella and Limnoecetis probably
share many similarities with the dipseudopsid larvae
known. The phylogeny also provides evidence for re-
moving Eodipseudopsis from the Dipseudopsidae (cf.
final section of text).
The sister group relationship of the genus
Phylocentropus remains uncertain. Phylocentropus phe-
netically agrees closely with Hyalopsyche, but the sim-
ilarities of these two genera are predominantly plesio-
morphic. Thus, the phylogenetic analysis above does
not provide supporting evidence for Hyalopsychidae
s. str. Schmid (1980) and in fact, infers that this tax-
on might be paraphyletic. Therefore we concur with
Ross and Gibbs for the most part on the classification
of Phylocentropus, and Wells and Cartwright on the
placement of Hyalopsyche and the suppression of the
Hyalopsychidae, a taxon that presently serves no use-
ful purpose in phylogenetics.
Keys to genera of Dipseudopsidae
Larvae
The larvae of Hyalopsychella Ulmer and Limnoecetis
Marlier are unknown.
1. Mandibles with apicolateral teeth; head capsule
short, with lateral margins nearly parallel and its
total length less than its maximum width (Ulmer
1957 GILES TO CS) EEEN LRE eens D
— Mandibles without apicolateral teeth; head cap-
sule long, with lateral margins tapering anteriad,
and with its length slightly greater than its maxi-
mum width (Wiggins 1977, Wells & Cartwright
LOOF N Ga EE TAR 3
2. Frontoclypeal suture V-shaped with lateral lines
tapering irregularly posteriad toward the dorsal
eoronallsurure Olmer 1957) er ne
RMN duo cetona Dipseudopsis Walker
— Frontoclypeal suture with the posterior portion
U-shaped, and the anterolateral lines nearly par-
allel between the eyes and then curving mesad to
meet the coronal suture posteriad (Gibbs 1968)
ennio dado Protodipseudopsis Ulmer
3. Frontoclypeal suture sessile in respect to posterior
margin, posterior 1/3 nearly regular V-shaped
and without hourglass-shaped constriction
(Wells & Cartwright 1993) .. Hyalopsyche Ulmer
— Frontoclypeal suture with short petiole joining
posterior margin, posterior 1/3 more irregular V-
shaped and with hourglass-shaped constriction
(Niseinsl97y) Phylocentropus Banks
Adults
1. Pronotum with setaceous warts conspicuous and
without deep median fissure … ne 2
— Pronotum with setaceous warts inconspicuous
and with deep median fissure … un 4
2. Labial palps present; forewing with fork I long
andasessiler a een Phylocentropus Banks
— Labial palps absent; forewing with fork I short
andipetiolateforzabsentg me... ehe 5
3. Fore and hind wings with fork I short and petio-
large anhand Hyalopsyche Ulmer
— Fore and hind wings with fork I absent ..............
tele NE Hyalopsychella Ulmer
4. Tibial spurs 1, 2, 2; female with maxillary palps
three-segmented; one species endemic to Lake
liansanyilkam sur. Limnoecetis Marlier
— Tibial spurs 3, 4, 4; female with maxillary palps
five-segmented; Africa and Asia...
5. Mouthparts with maxilla modified into a probos-
cis; hindwing with median cell closed; male hind
mesoapical tibial spur modified, often twisted
and/or branched; Asia and Africa …
EN mil heater. Les Sass, Jose Dipseudopsis Walker
— Mouthparts without a proboscis; hindwing with
median cell open; male hind mesoapical tibial
spur normal; Africa ...... Protodipseudopsis Ulmer
SYSTEMATIC PART
Dipseudopsis Walker, 1852
Dipseudopsis Walker, 1852: 91. — Type species: Dipseudopsis
capensis Walker (monobasic).
Nesopsyche McLachlan 1866: 168. — Type species: Neopsyche
flavisignata McLachlan (monobasic).
Esperona Navás 1915: 397. — Type species: Esperona orienta-
lis Navás (monobasic).
Bathytinodes Iwata 1927: 235. — Type species: Bathytinodes
albus Iwata (monobasic).
WEAVER & MALICKY: Dipseudopsis from Asia
Dipseudopsodes Lestage 1936: 170 (invalid, type species not
designated).
The genus Dipseudopsis was proposed by Walker
(1852) with the description of D. capensis from South
Africa and placed in the family Sericostomatidae
Stephens, a family designation which was common
for caddisflies, exhibiting bizarre characteristics, that
did not agree with the diagnoses of other families.
Later when McLachlan (1863) added a second species
to the genus, D. collaris from China, he placed Di-
pseudopsis in the family Rhyacophilidae. Brauer
(1868) placed it in the Hydropsychidae s. lat, near
Cyrnus Stephens. However, after examination of the
adult mouthparts, Ulmer (1904a, b) placed the genus
in the Polycentropodidae, subfamily Dipseudopsinae.
Accordingly, most workers have placed the genus in
Polycentropodidae (Navás 1913, Martynov 1935,
Schmid 1949). However, Banks (1931b) maintained
a conservative approach, having never recognized the
full family status of Polycentropodidae, and followed
Brauer’s placement of the genus in the Hydropsychi-
dae s. lat.
Etymology. — Dipseudopsis: Greek di, two; pseudos,
false; ops, eye; feminine. The actual etymology is un-
certain. The basis of this name is uncertain also, since
there are no noticeable characters such as wing spots
present in the type species of the genus, D. capensis.
Perhaps it was based on the appearance of the prono-
tum, having the setal areas expanded and divided by a
deep median fissure.
Adult
Head and thorax (figs. 12A-B, 13, 14): The head
has a sparse number of short inconspicuous setae and
is nearly glabrous; sometimes it is glossy and very gla-
brous. The setal areas of both the head and thorax are
not raised like typical setaceous warts, but are flush
with other surrounding sclerites. The setae of the bo-
dy and wings are mostly short sparse and inconspi-
cuous, giving the imago a glabrous appearance. The
pronotum is raised to the same level as the head and
mesonotum and is divided by a deep median furrow,
probably resulting from modifications of the lateral
setal areas, which have remained separate, but have
become both enlarged and glabrous. The tegula are
flat and sit like epaulets on the bases of the forewings.
Legs with tibial spur formula complete: 3, 4, 4. The
head, thorax, and sometimes the anterior segments of
the abdomen have patterns of yellowish or orangish
brown contrasting with dark brown. The forewing is
narrow, length ranging from 10 to 20 mm, usually
mostly brown with a hyaline spot at m-cu and the ar-
culus, and sometimes with a pattern of translucent
99
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
bands or spots against a dark background, or with a
striped pattern of dark veins against lighter translu-
cent membranes; fork I is either short and petiolate or
absent, II and IV are long and sessile, III and V are
long and petiolate, and the discal and medial cells are
closed. The hindwing is shorter and sometimes bro-
ader than the forewing, being somewhat triangular
and lighter in colour, sometimes nearly translucent;
forks II and V are long and sessile, forks I, III, IV ab-
sent, the discal cell closed, and unlike other dipseud-
opsid genera described the median cell is closed. The
scent glands of sternum V are present in the female
and absent in the male, their openings are rather in-
conspicuous and lack exterior projections. This con-
dition differs in Phylocentropus placidus (Banks) exa-
mined, having the scent gland absent in both sexes.
The mouthparts (fig. 12B, 13, 14) have been de-
scribed by Ulmer (1904a), Cummings (1913) and
Crichton (1957: fig. 53), and are quite unique, hav-
ing the lacinia modified into a proboscis, functioning
as a siphoning tube. The two processes which form
the proboscis are not held together in preserved spec-
imens, but curl away from each other distally. Each
process is shaped like half of a long slender straw, with
its longitudinal edges fringed with numerous short
scalelike setae, having many transverse ridges along its
shaft (thus being semiannulate), and having several
mesal dentations at its base. The proboscis is at least
as long as the first 2-4 segments of the maxillary
palps, or usually about the same length as the labium,
including base plus palps. The haustellum is slender,
about as long as the base of the labium, and its apex is
bilobed. The frontoclypeus is large and slightly bul-
bous, apparently well-developed to support a pump-
ing organ. Protodipseudopsis differs noticeably from
Dipseudopsis by lacking a well-developed proboscis
(fig. 12C); its frontoclypeus is shorter and less bul-
bous and apparently the pumping organ is reduced.
Male. — The male is dark brown, sometimes with
contrasting patterns in the wings and body, and more
striking than in the female. The mesoapical spur of
the hind tibia is modified, often twisted like a cork-
screw and sometimes branched into two or more pro-
cesses. It is interesting to note that the same spur is al-
so modified in Phylocentropus auriceps (Banks),
(Schuster & Hamilton 1984), being exceptional
among the other dipseudopsids.
Genitalia (fig. 17): The male genitalia are blunt,
generally lacking in conspicuous characteristics, and
the processes of the genitalia are retracted and ob-
scured by the large blunt preanal appendages. This
blunt appearance of the male terminalia, especially in
dry material, probably is why males sometimes have
been mistaken as females, e.g. the types of D. flavisig-
100
nata (McLachlan, 1866) and D. benardi Navás
(1930). Segment IX with articulation between tergite
and sternite, resembling an open jaw in lateral view;
the tergum and sternum also articulate with the pre-
anal appendages. The tergum of segment IX is much
smaller than the sternum and usually overhangs seg-
ment X, and its apical margin may be triangular,
rounded, or bilobed in dorsal view. The sternum of
segment IX is heavily sclerotized and is the most mas-
sive feature of genitalia, providing support for the in-
ferior appendages, the phallus, and the ventral articu-
lation of the preanal appendages. The sternum of
segment IX is a simple structure except for the meso-
superior processes that, along with segment X, form a
hood that hangs over the dorsal portion of the phallus
which is mostly membranous. Segment X is inferior
to segment IX, and subequal in length to the phallus.
The phallus is mostly membranous dorsad and heav-
ily sclerotized ventrad. Hence, segment X and the
phallus appear to function in concert, with segment X
providing dorsal coverage and support to the phallus.
The inferior appendages are thumblike, with apices
directly dorsad, perhaps modified to hold segment X
and the phallus together, as well as clasping the fe-
male during copulation.
Female. — Females are usually lighter in colour than
the males, and as a result they are more dull and have
fewer contrasting colour patterns in their wings and
bodies. However, the contrasting colouration of an
unidentified female specimen (fig. 2) may be excep-
tional. In several species, such as D. nervosa (figs. 3,
4), where the males have brown wings with large
translucent spots, the females have striped forewings
with dark veins against a translucent background.
Sometimes females have forewings with faded indis-
tinct spots that are remotely similar to translucent
spots in the forewings of males. However, the female
forewings are most often dull brown and show no
striking colour patterns.
Genitalia (figs. 15-16): Sternum VIII forms a sin-
gle plate, not divided into lateral lobes as in the
Polycentropodidae. Segment IX has a well-sclerotized
tergite, as in some species such as D. recta (fig. 15),
but is more membranous in others (fig. 16); the ster-
num is mostly membranous. The vaginal apparatus
and the vulval scale are attached to the sternum of
segment IX by sclerotized bands and membranous
folds. Both the vaginal apparatus and the vulval scale
are sclerotized. The length of the vaginal apparatus is
variable, being long in D. recta, where it extends ante-
riad reaching the anterior margin of sternite VIII (fig.
15), but is much shorter in other species (fig. 16).
The genitalia are generally similar to those of
Phylocentropus (Schmid 1980, Schuster and Hamil-
ton 1984), except that segment IX is more heavily
sclerotized. This diagnosis is provisional, because the
females of most species remain undescribed. We ob-
serve that the females of some species can be distin-
guished from others by colour patterns of the body
and wings, and characteristics of the genitalia.
Larva
The larva of Dipseudopsis was first described by
Iwata (1927), although under a different name. A key
separating the larvae of Dipseudopsis and Protodi-
pseudopsis was provided by Ulmer (1957), and a simi-
lar key was provided by Marlier (1962), whereby the
larvae of Dipseudopsis and Protodipseudopsis were dis-
tinguished by the gular sclerite, mandibles, and hind
tarsal claw. Protodipseudopsis and Dipseudopsis are
unique in having mandibles with apicolateral teeth, a
probable synapomorphy (Ulmer 1957: f. 401; Gibbs
1968: fig. 1; Marlier 1962: fig. 50; Marlier 1979: fig.
5). The known larvae of Dipseudopsis known can be
distinguished from those of other dipseudopsids by
having head capsule short, lateral margins nearly pa-
rallel with total length less than its maximum width,
frontoclypeal suture V-shaped and the lateral lines ta-
pering irregularly and posteriad toward the dorsal co-
ronal suture, and mandibles with apicolateral teeth.
Larval behaviour and habitat. — The larval tube-
dwelling and feeding-behaviour have been described
for Protodipseudopsis (Gibbs 1968) and Phylocentropus
(Wallace et al. 1976, Wiggins 1977). The larvae of
these genera live in the bottom substrates of streams
or lakes and construct a somewhat Y-shaped silken
tube composed of sand grains and detritus. The ends
of the two upper branches, usually one vertical and
the other inclined, are open and exposed to the
stream current, the rest of the tube is located within
the bottom substrate, and the end of the bottom
branch is closed. The opening of the vertical branch is
an intake and the inclined branch an outlet, through
which a directional flow is established via rapid oscil-
lations of the larva. This forced flow enables a fine
capture net, within an enlarged chamber of the outlet
branch, to collect fine organic food particles for the
larva to eat. The larval tubes of Hyalopsyche are
branched and appear to be similar to those of
Phylocentropus (Wells & Cartwright 1993), as are the
tube-dwellings of Dipseudopsis. Therefore, since
branched tube-dwellings are characteristic for the lar-
vae of Dipseudopsis, Hyalopsyche, Protodipseudopsis,
and Phylocentropus, this type of larval retreat and its
associated mode of feeding-behaviour is a synapo-
morphy of Dipseudopsidae. The larva retreat of the
hydropsychid genus Macrostemum Kolenati (Wiggins
1977, as Macronema Pictet) is generally similar, but
differs in many details and is certainly a homoplasy.
WEAVER & MALICky: Dipseudopsis from Asia
Ulmer’s (1957: 303) description of the larval re-
treat and biology of Dipseudopsis is translated as fol-
lows: ‘Larvae and pupae live in calm pools of rivers,
also in lakes and lake-outflows, in the mud or fine
sandy substrates; the larvae not free-living there, but
construct tubes out of secretions, covered with mud
etc., in the depths; these tube-dwellings are more or
less bent and twisted, elastic and flexible, but never-
theless fairly resistant; sometimes they are branched
into 2 or 3 tubes, but only one of the branches reach-
es the [substrate] surface and there it is closed, while
the other lower ends are open.’ It seems obvious that
Ulmer’s orientation of the tube-dwelling is upside
down; his description should depict the two open
ends reaching the surface, and the one closed branch
completely buried in the substrate. This would agree
more with the tube-dwellings described in detail for
Phylocentropus and Protodipseudopsis, and the descrip-
tion by Scott (1985: 334) for Dipseudopsis in south-
ern Africa: “The larvae are bottom dwellers in stand-
ing or slow-flowing water where they construct
U-shaped silk-lined tubes with a transverse net in
mud or silt or on submerged branches, filtering out
algae and organic particles on which they feed.’
The larvae and pupae of Dipseudopsis inhabit a
range of freshwater habitats, including the sandy dep-
ositional zones of lakes and the pool areas of rivers
and streams. Marlier (1953, 1962) reports that im-
matures in Africa were found in shallow water, while
Tsuda (1939) reported that larvae in Japan were dis-
covered at a depth of 90 m. The collections of D.
diehli Malicky & Weaver, at elevations of 1,400 m in
Sumatra, devoid of large rivers and lakes, suggests
that some species also live in mountain streams.
Distribution
Dipseudopsis is known to occur generally through-
out the old world tropics of Africa and Asia. The
African fauna recorded from the northern Nile River
Valley, including Egypt, Sudan, and throughout
most of the Afrotropical Region, from Ethiopia and
Kenya to Nigeria, Ghana and Sierra Leone, and to
Zaire, Mozambique, South Africa, and Madagascar.
The genus is represented by 42 described species, 23
from the African continent, and 19 from Madagascar.
However, in light of the problems discovered in this
work, we suspect that the African species are in need
of a thorough taxonomic revision. The Asian fauna
with 36 recognized species is known throughout most
of the Oriental Region and part of the southeastern
Palearctic Region, including Pakistan, India, Sri
Lanka, Nepal, Bangladesh, Burma, Thailand,
Cambodia, Vietnam, China, Japan, Philippines,
Malaysia, and Indonesia, including Sumatra, Java,
Borneo and Sulawesi.
101
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Checklist of Dipseudopsis from Asia
adiaturix sp. n. p. 102
benardi Navás, 1930. p. 104
langana Navás, 1930 syn. n.
arculata Navás, 1931 syn. n.
ulmeri Schmid & Denning, 1979 syn. n.
bicolorata Martynov, 1935. p. 107
collaris McLachlan, 1863. p. 109
stellata McLachlan, 1875 syn. n.
alba (Iwata, 1927).
bakeri Banks, 1916 syn. n.
discors Navas, 1924 syn. n.
contorta Banks, 1931b. p. 110
diehli Malicky & Weaver, 1988. paie
digitata Ulmer, 1907a. poll
doehleri Ulmer, 1929 (döhler:). p. 115
elongata Banks, 1920. p. 116
flavisignata (McLachlan, 1866) p. 116
flinti sp. n. p. 116
immaculata Ulmer, 1905. p. 118
moesta Banks, 1931b syn. n.
indica McLachlan, 1875. p. 120
buddha Banks, 1913 syn. n.
i. sindica Martynov, 1935 syn. n.
infuscata McLachlan, 1875. p. 120
knappi Schmid & Denning, 1979. pal?
lamellata Martynov, 1935. p. 121
lucasi sp. n. p. 123
maculata Ulmer, 1907a. pals
malaisei sp. n. p. 124
martynovi sp. n. p. 124
modesta Banks, 1911. p. 124
pallida Martynov, 1935 syn. n.
morosa Banks, 1924. 05.1125
nebulosa Albarda, 1881. p. 127
nervosa Brauer, 1868. 127
luctuosa Banks, 1913.
venosa Naväs, 1931 syn. n.
nervosella Ulmer, 1951 stat. n. p. 128
nieuwenhuist Ulmer, 1909. p. 130
notata (Fabricius, 1781). p. 130
horni Ulmer, 1915 syn. n.
recta Martynov, 1935. p. 131
bombayana Martynov, 1935 syn. n.
robustior robustior Ulmer, 1929 stat. n. p. 132
akhila Schmid & Denning, 1979 syn. n.
thailandica Schmid & Denning, 1979 syn. n.
junki Marlier, 1979 syn. n.
robustior andamanensis ssp. n. p. 134
schmidi sp. n. p. 134
spectabilis Banks, 1931a. p. 134
stabatensis Malicky & Weaver, 1988. 9.1135
tonkinensis Naväs, 1921. p- 135
triclavata Martynov, 1935. PAY
varians Ulmer, 1929. p. 138
102
petersorum Schmid & Denning, 1979 syn. n.
voluta Ulmer, 1906. p. 138
onychophora Navas, 1935 nomen dubium p.139
orientalis (Navas, 1913) nomen dubium p. 139
Dipseudopsis adiaturix sp. n.
(fig. 18)
Type material. — Holotype d : INDONESIA: Sumatra, Huta
Padang, 26.1.1990, Diehl (CLHM).
Male. — Head with dorsum and frontoclypeus dark
brown, almost black, but posterior dorsal warts yello-
wish brown. Basal segments of antennae brown (type
with distal parts of antennae and mouthparts missing
due to ant attack). Thorax, tergites and sternites dark
brown. Coxae of fore and mid legs brown, coxa of
hind legs yellowish brown. Basal % of fore femora
dark brown, femora of mid and hind legs with basal %
brownish; distal % of all femora, tibia of fore and mid
legs, and all tarsi yellow; distal 4 of hind tibia brown.
Modified spur (fig. 18) long and slender, more than %
as long as adjacent tarsal segment, more than 2x long-
er than outer spur, distal claw about % as long as spur
and bent obliquely laterally, and with short lateral
thorn at # length from base. Forewing 16 mm, slen-
der, dark brown, but with nervation darker, having
hyaline lunula at m-cu, and indistinct translucent
spots between M, and Cu, and near apex of 1A and
2A. Hindwing 10 mm, dark brown, with hyaline lu-
nula at m-cu. Abdomen yellowish brown. Genitalia
(18A-D): Tergum IX broad and bilobate with me-
dian sized cavity below in dorsal view; sternum IX
with mesosuperior process fingerlike in lateral view.
Dorsal edge of segment X with strong incision in lat-
eral view, but apex not incised. Preanal appendages
long and triangular in lateral view. Inferior appenda-
ges curved dorsad in lateral view, with distal part
slightly clavate and basolateral mesal margins squarish
in caudal view. Phallus calyx-like in ventral view,
about % as long as segment X.
Etymology. — A noun in apposition: named after a
Celt who lived in Austria 2000 years ago.
Distribution. — Indonesia: Sumatra.
Remarks. — This species is only known from the
unique type. It has a modified spur similar to D. flav-
isignata, being long slender, acuminate, slightly sinu-
ate and bearing a short subapical lateral thorn.
However, it differs in having the male modified spur
with the lateral thorn closer to its apex, rather than
near the base, the male forewings mostly dark brown
and without a striking pattern of translucent stripes
and by characteristics of the male genitalia.
Pictorial atlas (p. 103-106)
A pictorial atlas is provided for quick species deter-
mination. The figures of the following male charac-
ters are provided for each species: 1) dorsal aspect of
segment IX and X, 2) shape and relative size of the left
modified spur, and 3) caudal aspect of the right infe-
D. nebulosa Albarda
A |]
Burma, Thailand,
Sumatra, Malay Pen. vA
D. immaculata Ulmer
CAN?
Sumatra, Borneo, Malay Pen.
D. schmidi Weaver and DI
India, Bangladesh
D. varians Ulmer
1
Thailand, Malay Pen. il
D. doehleri Ulmer i
SW
Assam, Burma, Thailand
WEAVER & MALICKY: Dipseudopsis from Asia
rior appendage (but for subspecies, only figures of the
modified spur). The distributional data are listed be-
low the figures. Possible identifications should be
confirmed with the complete species descriptions and
figures in the text.
D. flavisignata (McLachlan)
Be
Sulawesi
D. adiaturix Weaver
and Malicky
>
Sumatra
D. voluta Ulmer
Dr
Sulawesi
D. lamellata Martynov
India, Burma, Sip
D. indica SY \
India, Bangladesh
103
PF |”
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
D. infuscata McLachlan
Ss
Sumatra, Borneo, Java, Sulawesi
D. collaris McLachlan
Ny
China, Japan, Philippines
D. robustior Ulmer
ANY
Burma, Thailand, Vietnam,
Cambodia, Malay Pen.
D. r. andamanensis
Weaver and Malicky
Andaman Is.
D. notata (Fab.)
India, Sri Lanka
Dipseudopsis benardi Navás, 1930
(figs. 10, 19)
Dipseudopsis stellata. — Ulmer 1905: 96, fig. 75b [misdet].
Dipseudopsis benardi Navás, 1930: 141, Type d *, VIETNAM
‘Tonkin’: Phu Lang Thuong, 1909, det. P. Navás S. J.,
G. Benard (MNHN); 1d*, same data (MNHN). — Fischer
104
D. diehli Malicky and Weaver
Sumatra \
D. spectabilis Banks
Pri
Borneo
D. knappi Schmid and Denning
AAW
Thailand
D. nervosa Brauer
Zij
Philippines
wij
1962: 5; Fischer 1972: 3.
Dipseudopsis langana Navás, 1930: 141, Type 9*, VIETNAM
‘Tonkin’: Phu Lang Thuong, 1909, det. P. Navás S. J., G.
Benard (MNHN). — Fischer 1962: 10; Fischer 1972: 5. Syn. n.
Dipseudopsis arculata Navás, 1931: 85-86, Holotype d*,
CHINA: Guangfong: ‘Cong-tcheou Canton (MZBS). —
Fischer 1962: 5; Fischer 1972: 2. Syn. n.
D. morosa Banks
Me
Philippines W
D. nervosella Ulmer
AWA
Borneo
D. maculata Ulmer
>
En) à
D. nieuwenhuisi
ane
D. digitata Ulmer
avs
Philippines
Dipseudopsis moesta. — Ulmer 1951: 128 [misdet].
Dipseudopsis ulmeri Schmid & Denning, 1979: 243, fig. la-
e, Holotype d*, THAILAND: Chiang Mai Prov., E fork
Mae Ping, 56km N of Chiang Mai, 1300ft, at light,
24.X1.1964, Peters (CLDD). Syn. n.
Specimens examined. — CAMBODIA: 1d 29, ‘Cambodge’
WEAVER & MALICKY: Dipseudopsis from Asia
D. bicolorata Martynov
AW 0
India
D. contorta Banks
AMI
Sulawesi, Borneo, Java, Malay Pen.
D. martynovi Weaver and Malicky
i \ =
Philippines
D. tonkinensis Navas
ANGE
Vietnam, Hainan
Erratum: D. contorta, read Sumatra for Sulawesi.
Pavie, 1886 (MNHN). — CHINA: Fujian: 1d, ‘Foochow
Fuchou, 1936-37, det. D. stellata by Mosely, M. S. Yang (-
BMNH). — THAILAND: 26 , Chiang Mai, 19.V.1952, D. &E.
Thurman (USNM). 1d, Doi Sutep, 1952, D. & E. Thurman
(MCZC). 16, Bangkok, 5.X.1929, H. Smith (MCZC). 1d,
Ping River, Chiang Mai, 9.1V.1989, Malicky (CLHM). 26,
Ping River near Lampun, 20.X11.1989, Malicky (CLHM).
105
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
D. stabatensis Malicky and Weaver
yd
Sumatra, Borneo
D. flinti Weaver and Malicky
3
Selangor
D. malaisei Weaver and Malicky
AWC
D. benardi Navas
Thailand, Vietnam, Cambodia, China
India, Nepal
Id, Chiang Mai Zoo, light trap, 24-31.X.1988,
Chantaramongkol & Malicky (CLHM). 26, Kanchanaburi,
Sai Yok, 400ft, 4.V.1990; 66, 14.V.1988; Allen (CLHM).
2d, Kwei river, 11.X.1979 (CLHM).
Male. — Head dorsum mostly dark brown but pos-
106
1" I D D. A Banks
D. recta Martynov a
D. modesta Banks
AM
D. triclavata Martynov ,
SW
India, Sri Lanka
D. lucasi Weaver and Malicky
Wb
le
terior warts light brown; frontoclypeus light brown
but dorsal portion darker; antennal scape and pedicel
light brown, flagellum dark. Pronotum, mesonotum
and tegula dark brown. Forewing (fig. 10): 13.5-16.5
mm, mostly dark brown, but with translucent spots:
distinct small translucent spot just distal of r; small
ar
India
Figures 1-2. — Fig. 1, Dipseudopsis bicolorata Martynov,
male, dorsal. Fig. 2, Dipseudopsis sp. female, dorsal, India,
Manipur, Nungha.
faded spots just distal of transverse cord, hyaline lu-
nula at m-cu, large translucent spot at arculus. Legs
brown. Modified spur (fig. 19): basal % unmodified
and apical % bifid with two short acuminate points
curved, spiraling opposite each other distally, each
point slightly longer than width of main shaft.
Genitalia (fig. 19): Tergum IX with posterior margin
curved in dorsal view, with posterior % extending over
segment X in lateral view; sternum IX having mesos-
uperior process with short dorsal triangular point and
long slender distal point, extending above basal % of
phallus in lateral view. Segment X tapering distally
into slender lobe in lateral view, extending posterior-
ly distal to preanal appendages about as far as inferior
appendages, apex bilobed. Preanal appendages with
posterior margin not distinctly incised in lateral view,
having blunt dorsal point, dorsal and ventral margins
converging to truncate posterior margin. Inferior ap-
pendages thumblike with obtuse midventral angle in
lateral view.
Distribution. — Cambodia; China: Guangdong,
Fujian; Thailand; Vietnam.
Remarks. — This species can be recognized by hav-
ing the modified spur with two short apical points
WEAVER & MALICKY: Dipseudopsis from Asia
spiraling opposite from each other, and the forewing
longer and darker brown than most other species in
southern China and Indochina. The types of D. be-
nardi, D. arculata, D. langana, and D. ulmeri were ex-
amined, and the latter three are recognized as junior
synonyms of D. benardi. The type of D. benardi is a
male, not a female as previously reported by Navas
(1930). The type of D. langana is recognized as a fe-
male of D. benardi, because it is has an identical loca-
lity label as the type of D. benardi and is similar in size
and general appearance.
Ulmer (1905: fig 75b) provided a figure of the
male modified spur of an undetermined specimen of
Dipseudopsis from ‘Cambodscha, Museum Paris’ and
Ulmer (1951: 128) later identified this figure as D.
moesta (syn. n. of D. immaculata). Ulmer’s figure is
more similar to the spur of D. benardi, and is prob-
ably not D. immaculata because this species is un-
known from Cambodia. Also, we have examined a
male specimen of D. benardi from Cambodia in the
MNHN, which might be the same specimen figured by
Ulmer.
Dipseudopsis bicolorata Martynov, 1935
(figs. 1, 20)
Dipseudopsis bicolorata Martynov, 1935: 165-166, figs. 68a-
c, 69a-b, Type series: INDIA: ‘26, 2. Castle Rock, North
Kanara district, Bombay Pres. October 1916. S. W.
Kemp … 6. Talewadi, Castle Rock, N. Kanara district,
Bombay Pres. 3-10.1916. S. W. Kemp’. The type materi-
al present in the collection of the Zoological Survey of
India is in poor condition and too delicate to send by mail
(pers. comm. Dr. K. Rai). Syntype 19 *: det. Martynov,
Karnataka, N. Kanara dist., Castle Rock, in alcohol,
X.1916, S. Kemp (ZRAS). — Fischer 1962: 5; Higler 1992:
80.
Specimens examined. — INDIA: Karnataka: 16, Shimoga
Dist, Agumbe Ghat 2000ft, V.1974, Lucas (CLHM). Tamil
Nadu: 4d, Kattalaimala, 25-26.XII.1961, F. Schmid (-
CNCI). 16, Ottakada, 5.1.1962, F. Schmid (CNCI).
Male. — Body and wings (fig. 1). Head dorsum and
pronotum orangish brown, frontoclypeus mostly
dark brown but dorsal margin orangish brown, genae
yellowish, antennal scape orangish, pedicel and flagel-
lum brown, palps dark brown, but stipes orangish
brown, maxillary palps 3.2 mm, proboscis 1.7 mm, as
long as first three segments of maxillary palps, labial
palps 1.4 mm; cervical sclerites and mesonotum dark
brown. Forewing 16-21 mm, almost unicoloured
dark brown, but darker along anterior margin
between C and R and posterior margin, only hyaline
spots at m-cu and arculus. Front leg brown, middle
and hind legs with coxae, trochanters and femora
orangish brown, distal segments brown. Modified
spur (fig. 20) having one apical point, curving in
107
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
li
ae 7
TE 7/55
e
7
Mi Ye di
7,
Figures 3-11. — Fig. 3-4, Dipseudopsis nervosa Brauer: Fig. 3, male. Fig. 4, female. Fig. 5, D. doehleri Ulmer, male. Fig. 6, D.
nieuwenhuisi Ulmer, male. Fig. 7, D. nebulosa Albarda, male. Fig. 8, D. digitata Ulmer, male. Fig. 9, D. spectabilis Banks, male
wings. Fig. 10, D. benardi Navás, male forewing. Fig. 11, D. flinti, n. sp., male forewing.
108
Figures 12A-C. — Fig. 12A,
Dipseudopsis nebulosa Al-
barda, male, head and tho-
rax, dorsal. Fig. 12B, D.
doehleri Ulmer, male, head
with mouthparts, ventral.
Fig. 12C, Protodipseudopsis
sjoestedti Ulmer, male, head
with mouthparts, anterior.
plane perpendicular to longitudinal axis of spur.
Genitalia (fig. 20): Tergum IX triangular with
blunt apex in dorsal view, posterior margin only
slightly overhanging segment X in lateral view.
Segment X ellipsoidal with length 14 X basal width
in dorsal view, tapered distad and apex incised.
Preanal appendages with dorsal margin irregular in
lateral view, directed dorsad with apex obtuse or
sometimes rounded and ventral margin broadly
rounded. Inferior appendages with massive dorsal ex-
tension that doubles height at middle as compared to
base in lateral view, with distinct apicomesal tooth in
ventral view, and with curved apicolateral flanges.
Distribution. — India: Karnataka, Tamil Nadu.
Remarks. — This is the largest species of Dipseudo-
psis known from India and is easily recognized by its
colouration of the males which superficially resemble
ctenuchid moths. The orangish brown dorsum of the
head and pronotum contrast strikingly with the uni-
coloured, dark brown mesonotum and forewings.
The modified spur bears a single curved apical point
that lies in a plane perpendicular to the longitudinal
axis of the spur. The specimen in fig. 2 is possibly a
female of this species.
WEAVER & MALICKY: Dipseudopsis from Asia
Dipseudopsis collaris McLachlan, 1863
(fig. 21)
Dipseudopsis collaris McLachlan, 1863: 496, pl. 19: fig. 6,
Lectotype d *, selected by Kimmins (1957: 101), Hong
Kong: (BMNH). Cotype 1d*: Hong Kong: Soochow
(MCZC 11080). Weidner (1964): Paratype 2, CHINA:
Hong Kong (ZMUH). — Fischer 1962: 7; Weidner 1964:
73; Fischer 1972: 3.
Dipseudopsis stellata McLachlan, 1875: 16-17, pl. 2: fig. 11, Ho-
lotype d (Kimmins, 1957: 101): [CHINA: Zhejiang:] Shang-
hai, McL. coll. (BMNH); 38* [possible syntypes]: Shanghai,
McL. coll. (BMNH); ‘Paratype’ 19*: ‘N China’ McL. coll
(BMNH). — Fischer 1962: 15. Fischer 1972: 7-8. Syn. n.
nec Dipseudopsis stellata. — Ulmer 1915: 73 (Sri Lanka);
Lestage 1925: 65, 115 (Java); Banks 1931b: 401 (stellatus,
Thailand); Tsuda 1942: 266 (Thailand); Schmid 1958:
35, 89 (Sri Lanka).
Dipseudopsis nebulosa. — Banks 1914: 178 (Luzon) [misdet.].
Dipseudopsis bakeri Banks, 1916: 215-216, pl. 2: figs. 20,
21, Type d *, PHILIPPINES: Luzon, Laguna, Mt Maquiling
and Los Baños,Baker (MCZC 11768). Only 1d at the
MCZC labelled as type, although several other specimens,
not labelled as types, at the MCZC and USNM are part of
the type series. — Fischer 1962: 5; Fischer 1972: 3. Syn. n.
Dipseudopsis discors Navas, 1924: 205. — Fischer 1962: 8.
Syn. n.
Dipseudopsis morosa. — Banks 1924: 450 (pro parte). Syntype
16 * is actually D. collaris. Luzon: Mt Maquiling, Baker
(MCZC 14826).
Bathytinodes alba Iwata, 1927: 209-210, pl. 4: fig. 87-91.
Dipseudopsis alba (Iwata, 1927). — Fischer 1962: 217;
Fischer 1972: 2.
109
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
wo; frontoclypeus
tentorial pit
labrum
lacinia
haustellum
labial palp maxillary palp
13
Figures 13-14. — Fig. 13, Dipseudopsis sp., male head, anteri-
or, modified after (Crichton 1957). Fig. 14, Dipseudopsis
doehleri Ulmer, male head and prothorax, lateral.
Specimens examined. — CHINA: Guangdong: 3d,
Hushan, 3.VI.1963, Tian (CLHM). Jiangxi: 1d 2d, det.
Ulmer (1905) as D. stellata, ‘Kiang-si’, 1869, A. David
(MNHN). Hong Kong: 1d, Foochow (MCZC). Jiangsu: 64,
Fulin, 18.VIIL1987, Tian (CLHM). 86 1%, Jinhu,
22.VI.1987, Sun Changhai (USNM). Zhejiang: 16 19,
‘Chekiang Hangtchéou’, Hangzhou, 1925, A. Pichon
(MNHN). 24 19, Shanghai, det. McLachlan as D. stellata,
(BMNH). 24, Shuian, 20.VI.1957, Tian (CLHM). JAPAN: 16,
Lewis, McL. coll (BMNH). PHILIPPINES: Luzon: 88 19 Los
Bafios (MCZC), 276 (USNM), 16 (BMNH). 16, Manila
(MCZC). 336 69, Mt Maquiling, Baker (USNM), 26
(ENCI), 76 42 (MCZC), 1d 16 (BMNH). 34 19, Balinag,
A. P. Ashby (USNM).
Male. — Head, thorax, wings and legs with colour-
ation similar to D. robustior. Forewing 13-15 mm,
with circular pattern of 5-7 small translucent spots
110
around discal cell, having most spots distal to cord,
and spot at arculus adjoining hyaline area at apex of
Cu,. Modified spur (fig. 21) apical % bifid, crescent-
shaped, with two subequal curved points. Genitalia
(fig. 21): Tergum IX triangular in dorsal view, slight-
ly overhanging segment X in lateral view; sternum IX
mesosuperior process ellipsoidal with blunt apex
pointed posteriad, extending above basal % of phallus.
Preanal appendages with posterior margin slightly in-
cised in lateral view, dorsal margin at right angle to
posterior margin, ventral margin at obtuse angle,
sometimes varied with ventral margin continuous
with posterior margin. Inferior appendages fingerlike
with short basoventral shelf in lateral view.
Distribution. — China: Guangdong, Hong Kong,
Jiangsu, Jiangxi, Zhejiang; Philippines: Luzon; Japan:
Honshu.
Remarks. — This species is similar to D. robustior,
but differs by having inferior appendages more broad
and fingerlike in lateral view (although this is vari-
able) and modified spur with two apical points both
crescent-shaped and subequal in length. In D. robust-
ior the inferior appendage is more slender and angled,
its modified spur bears two points, one point slender-
er, more curved and more than 2x longer than other.
Previous records of D. collaris, as D. stellata, from Sri
Lanka, Java and Thailand are probably in error.
Dipseudopsis contorta Banks, 1931
(fig. 22)
Dipseudopsis sp. — Ulmer, 1905: 96, fig. 75c (not fig. 75e as
listed by Fischer 1962).
Dipseudopsis infuscata. — Ulmer 1905: 96, fig. 75d; Ulmer
1951: 128 [prob. misdet.].
Dipseudopsis contorta Banks, 1931b: 400-401, fig. 20, lecto-
type by present designation d*: MALAYSIA: West Ma-
laysia: Selangor, Kuala Lumpur, 29.1.1924, at light, H.
M. Pendlebury (BMNH); paralectotype 1 d *, without hind
legs: Pahang, Kuala Tahan, 300ft at light, 23.X1.1921, H.
M. Pendlebury (MCZC 16480). — Fischer 1962: 8;
Malicky & Weaver 1988: 3-4, fig. 3a-e.
Specimens examined. — 26 , ‘Borneo’ det. Ulmer 1905 as
D. infuscata (MNHN). INDONESIA: Kalimantan: 1d,
Pontianak, VII.1907, F. Muir (USNM). Java: 14, [Muller?]
(RMNH). Sumatra: 1d, Dolok Merangir, 19.VII-20.VI-
II.1971, Diehl (CLHM). 16, same data, V-X.1980, Diehl
(CLHM). 1d, Ost-Aceh 26-30.VIII. 1972, Diehl (CLHM).
16, NE Sumatra, 16.1-18.11.1979, Diehl (CLHM).
Male. — Head dorsum dark brown, except posteri-
or warts orangish brown, frontoclypeus dark brown,
antennae orangish brown; cervical sclerites dark
brown, pronotum light orangish brown and mesono-
tum dark brown. Maxillary palps 2.2 mm, brown;
Figures 15-16. — Fig. 15,
Dipseudopsis recta Marty-
nov, female: 15A, genitalia,
lateral; 15B, genitalia, ven-
tral; 15C, abdominal seg-
ment VII, lateral. Fig. 16,
Dipseudopsis sp. female,
‘Manipur Nungha’ (CNC):
16A, genitalia, lateral; 16B,
genitalia, ventral.
416A
proboscis 1.6 mm, as long as first two segments of
maxillary palps. Forewing 10-12.5 mm, brown with
row of lighter spots from stigma to Cu, just distal to
cord, and large translucent spot at arculus, hyaline
spot at m-cu. Legs mostly light brown, except front
and middle coxae dark brown. Modified spur (fig.
22) distal 4 branched into two subequal curved
points, with points diverging basally but converging
distally, sometimes with rounded subapical shelf.
Genitalia (fig. 22): Tergum IX semicircular in dorsal
view, with posterior margin having almost no exte-
nsion overhanging segment X in lateral view; sternum
IX with mesosuperior process short, slender, acumi-
nate having apex curved dorsad in lateral view.
Segment X ellipsoidal in dorsal view, long trapezoidal
in lateral view, extended distad beyond apex of inferi-
WEAVER & MALICKY: Dipseudopsis from Asia
or appendage. Preanal appendages variable, either D-
shaped with lobe extending distally as far as inferior
appendages in lateral view, or with posterior margin
incised and with short dorsoapical blunt point.
Inferior appendages with basoventral notch and apic-
oventral obtuse angle in lateral view, apical % variable,
either equal in height to base and truncate, or some-
times more expanded and clavate; with apex and apic-
olateral margin curved in ventral view, but bearing
subapicomesal obtuse tooth.
Distribution. — Indonesia: Kalimantan, Sumatra,
Java; Malaysia: West Malaysia.
Remarks. — This species is similar to D. collaris, but
differs by having the inferior appendage capitate, with
111
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
tergite IX
preanal appendage
So e
HE PTE
È AT
seo x
: ETS .
pr on
2
in Aa segment X
cia,
inferior appendage
sternite IX
17
Figure 17. — Fig. 17, Dipseudopsis sp., male genitalia, lateral.
a basodorsal notch and an apicoventral obtuse angle
in lateral view, and the modified spur with apical
points directed more laterally. Slight variation is no-
ticeable in the male modified spur and genitalia (cf.
Malicky & Weaver 1988).
Dipseudopsis diehli Malicky & Weaver, 1988
(fig. 23)
Dipseudopsis diehli Malicky & Weaver, 1988: 8, fig. Ga-f.
Holotype d*: INDONESIA: Sumatra: Prapat, 1400m,
Diehl (CLHM); paratypes 27 d : Prapat, Bukit Tinggi, Aek
Tarum, Sitahoan, Dolok Merangir (CLHM, CLJW).
Specimens examined. — INDONESIA: Sumatra: Pematang
Siantar: 54, 1.VII.84-13.IV.1985; 36, 1.VI-12.XII.1985;
36, 22.JV-11.VII.1986; 56 , 1.IX.88-15.1.1989; 15, 9.III-
10.X.1989, Diehl (CLHM).
Male. — Head dorsum yellowish brown with dark
semicircular spot at anterior margin. Frontoclypeus
brown. Scape yellowish brown, distal portion of an-
tenna brown. Maxillary palps 3.0 mm, proboscis 2.4
mm, extending as far as 44 segment of maxillary palp.
Cervical sclerites and pronotum yellowish brown.
Mesonotum and tegulae brown, with scutellum yello-
wish brown. Forewing 16-20 mm, brown with four
translucent areas: 1) long stripe adjacent to costal
margin; 2) ellipsoidal spot between stems of R and M;
3) transverse band distal to cord, between R, and M,
with two V-shaped incisions along distal margin, and
having narrow connection with anterior margin; 4)
triangular spot at arculus. Hindwing unicoloured
brown, but with hyaline comma at m-cu. Legs yello-
wish brown, with light brownish tibiae, apparently
112
caused by more dense covering of fine brown setae.
First segment of hind tarsi uniquely bent mesad
about % of its length to permit free motion of broad
modified spur. Modified spur (fig. 23) with apex bi-
fid with two sinuate points nearly parallel and slight-
ly unequal in length, both curved mesad, but apices
curved distad. Genitalia (fig. 23): tergum IX with ap-
ical 4 hanging over base of segment X in lateral view;
broad and short with posterior margin bilobed or
slightly triangular in dorsal view, anterior margin
slightly bilobed. Sternum IX mesosuperior process
extending to end of phallus. Segment X ellipsoidal in
dorsal view. Preanal appendages triangular in lateral
view with dorsal margin slightly inclined dorsad and
ventroposterior margin curved steeply dorsad.
Inferior appendages fingerlike and with ventral mar-
gin slightly concave in lateral view, but dorsal margin
serrate in middle; with U-shaped basomesal notch in
ventral view, distal arms slightly diverging apically
and each tapering into small apical lobe.
Distribution. — Indonesia: Sumatra.
Remarks. — This species is easily recognized by hav-
ing the male modified spur extremely broad and bi-
furcate, with points curved a wide whorl, and by hay-
ing forewing with large translucent spots. Some
specimens are darker brown, having brown parts that
are usually yellowish brown in other species.
Dipseudopsis digitata Ulmer, 1907
(figs. 8, 24)
Dipseudopsis digitata Ulmer, 1907a: 39-40, figs. 56-57,
Holotype d *: MALAYSIA: Banguey Is, Staudinger (ZMHB).
— Fischer 1962: 8; Fischer 1972: 3.
Specimens examined. — PHILIPPINES: Balabac: 26,
Dalawan Bay, 5.X.1961; 1d, 9.X.1961; 3d, 12.X.1961
(ZMUC). Busuanga: 5d, 4 km N San Nicolas, 20-
29.V.1962, H. Holtmann (BPBM). Culion: 66 19, 6km W
Culion, 7-11.VI.1962, light trap, H. Holtmann (BPBM).
Palawan: 1d, Brooke’s Point, Macagua, 75m, 1-4.1V.1962
(BPBM). 196 19, Tarumpitao Point, H. E. Milliron (BPBM).
176 72, Irawan, 14km W Puerto Princessa, 5m, D. R.
Davis (USNM). 1d, Litso, Amoyan Ck., 57km N Puerto
Princessa, 10m, 10.XII.1965, D. R. Davis (USNM). 1d,
Mainit, 11km NW Brooke’s Point, 18.XI.1965, D. R.
Davis (USNM). 1d, Chromite Mine, 28km W Puerto
Princessa, 400m, 1-7.XII.1965, D. R. Davis (USNM). 306,
Brooke’s Point, Uring Uring, 14-23.VIII.1961 (ZMUC).
Male. — Body and wings (fig. 8). Head, thorax and
forewing with pattern of light and dark bands. Head
dorsum with dark brown median band widening an-
teriad and narrowing posteriad, lateral areas light
brown and posterior warts lighter. Frontoclypeus
mostly dark brown, but with yellowish V-shaped
band pointed dorsad. Antennae light brown.
Maxillary palps 2.3 mm, proboscis 1.4 mm, as long as
WEAVER & MALICKy: Dipseudopsis from Asia
“
»
4
Ne
NÉ
aa)
’
pep
L
oa |
1
‘
?
\
A
.
SEN
>. L,
Serum"!
Figures 18-21. — Dipseudopsis, A-H, male genitalia: A, lateral; B, preanal appendages, segments IX and X, dorsal; €, inferior
appendages, ventral; D, phallus, lateral; E, phallus, ventral; F, segments IX, X, dorsal; G, ventral; H, left inferior appendage,
ventral. I-J, male modified apical spurs of hind tibia, ventral. Fig. 18, D. adiaturix, n. sp. Fig. 19, D. benardi Navas. Fig. 20,
D. bicolorata Martynov. Fig. 21, D. collaris McLachlan, 21H, type of D. stellata McLachlan.
113
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figures 22-25. — Dipseudopsis, A-H, male genitalia: A, lateral; B, preanal appendages, segments IX and X, dorsal; C, inferior
appendages, ventral; D, phallus, lateral; E, phallus, ventral; F, segments IX, X, dorsal; G, ventral; H, segment IX, dorsal. I-J,
male modified apical spurs of hind tibia, ventral. Fig. 22, D. contorta Banks. Fig. 23, D. diehli Malicky and Weaver. Fig. 24,
D. digitata Ulmer. Fig. 25, D. doehleri Ulmer.
114
first three segments of maxillary palp. Cervical scler-
ites dark brown ventrally, light brown dorsally.
Pronotum with raised areas light brown, median fur-
row and lateral sides dark brown. Mesonotum and te-
gula dark brown. Forewing 10-14 mm, with striped
pattern of dark brown veins and lighter cells, notice-
ably darker along thicker veins, R, and Cu,. Legs with
coxae dark brown and trochanters and distal seg-
ments light brown, except front coxa bicoloured, dark
brown ventrad and lighter dorsad. Modified spur (fig.
24) with three minute apical points and one long acu-
minate subapical point curved laterally, all points dis-
tal to blunt subapical shelf. Genitalia (fig. 24):
Tergum IX short and truncate in lateral view, posteri-
or margin not overhanging segment X, rectangular in
dorsal view; sternum IX small, with massive mesos-
uperior process having large broad dorsal lobe and
slender posteroventral process extending above basal
4 of phallus. Segment X with pair of minute basodor-
sal lateral lobes, dorsal margin angled obliquely ven-
trad in lateral view, but with apex turned dorsad, ven-
tral margin straight but also turned dorsad apically;
about % as wide as tergum IX and about twice as long
as wide in dorsal view, lateral sides parallel but ta-
pered into triangular apex, and apex slightly incised.
Preanal appendages broad and not incised in lateral
view, somewhat triangular with dorsal margin and
ventral margin curved and converging at blunt dorsal
apex. Inferior appendages capitate in lateral view,
dorsal margin concave, ventral margin straight with
short basal shelf, apex truncate with corners slightly
curved, extended dorsad.
Distribution. — Malaysia: Banguey Is.; Philippines:
Balabac, Busuanga, Culion, Palawan.
Remarks. — This species is easily recognized, having
the male segment X with a pair of minute dorsolateral
lobes at its base, and modified spur with three minute
apical points, one long acuminate subapical point
curved laterally, and all points distal to a squarish sub-
apical shelf.
Dipseudopsis doehleri Ulmer, 1929
(figs. 5, 12B, 14, 25)
Dipseudopsis döhleri Ulmer, 1929: 189-190, fig. 27,
Holotype d *: INDIA: Meghalaya: Khasi Hills, McL. coll
(BMNH). — Fischer 1962: 8.
Dipseudopsis doehleri Ulmer. — Fischer 1972: 4; Higler 1992:
80
Specimens examined. — BURMA: 14, Washaung, 20km E
of Myitkyina, 200m, 4.VII.34, Malaise (NHRS). 16,
Myitkyina, 175m, 1934, Malaise (BMNH). INDIA: Assam:
1d, 8mi E Ledo, 125m, 27.34N 95.34E, 13.X.1961, Ross
& Cavagnaro (CASC). 29, Abhoypur For., Naphuk, 360m,
12.X.1961, Ross & Cavagnaro (CASC). THAILAND: 9d,
WEAVER & MALICKY: Dipseudopsis from Asia
Nakhon Nayok Prov., Khao Yai National Park, 700m,
29.IX-6.X.1984, Karsholt, Lomholdt & Nielsen (ZMUC).
73, Nam Nao, Pet Cha Boon, 5.V.1989, Allen (CLHM).
208, Puk Hieo, Chayapoom, 6.V.1989, Allen (CLHM).
Male. — Body and wings (fig. 5). Head and all of its
appendages dark brown, except proboscis yellowish.
Mouthparts (fig. 12B, 14): Maxillary palps 3.6 mm;
proboscis 2.7 mm, extended to midlength of apical
segment of maxillary palp; haustellum 0.8 mm; labial
palps 1.7 mm. Cervical sclerites and prothorax ivory.
Mesonotum, tegula and metanotum dark brown.
Forewing 16-17 mm, dark brown with three distinct
translucent spots: 1) basal triangle between R, and
Cu, from base to fork I of Rs, 2) subapical spot distal
to cord with apical margin incised at forks II, and III)
ellipsoidal spot at arculus, similar as in D. nebulosa ex-
cept for translucent spot at arculus. Legs dark brown,
except front coxa with lateral side ivory. Hindwing
with large translucent triangle as in front wing, apical
region brown, anal region translucent, Cu, dark
brown. Modified spur (fig. 25) similar to D. immacu-
lata, unbranched, slightly longer than adjacent spur,
basal 4 unmodified, apical 4 twisted 180° into short
spiral, apical point directed mesad. Abdomen with
anterior sternites II-V ivory, giving translucent trian-
gles of forewings a white background so that contrast-
ing pattern is displayed when wings are folded, ster-
nites VI-VIII dark brown. Genitalia (fig. 25):
Tergum IX triangular with posterior margin rounded
in dorsal view, apical % overhanging segment X in lat-
eral view. Segment X long slender in lateral view, with
short base having parallel sides in dorsal view, and ap-
ical portion cordate and apex deeply incised. Preanal
appendages abbreviated with slender dorsoapical
point in lateral view, but truncate and triangular in
dorsal view. Inferior appendages long and thumblike,
without incisions or points, curved slightly dorsad in
lateral view.
Distribution. — Burma; India: Assam, Meghalaya;
Thailand.
Remarks. — This species is most similar to D. neb-
ulosa, but differs by having the head entirely dark
brown and pronotum ivory, forewing with translu-
cent round spot at the arculus, and modified spur rel-
atively shorter, with less than % of apical portion
twisted. It also differs by having male genitalia with
segment X and the preanal appendages more slender
in lateral view, and segment X also bears a unique pair
of basolateral humps in dorsal view. A teneral male
specimen from Hainan Island, possibly an unde-
scribed species near D. doehleri, was observed in the
collection at the MCZC.
1115
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Dipseudopsis elongata Banks, 1920
(fig. 26)
Dipseudopsis elongata Banks, 1920: 361-362, pl. 1: fig. 13,
Type d*: MALAYSIA: Sabah, Sandakan, Baker (MCZC
10882). — Fischer 1962: 8-9; Fischer 1972: 4.
Specimens examined. — MALAYSIA: Sabah: 14, Sandakan
Bay, NW Sepilok For. Res., 1-10m, 26.X.1957, J. L.
Gressitt (BPBM). 1d, ‘North Borneo’ McL. coll (BMNH).
Male. — Head dorsum glossy dark brown, but pos-
terior warts light brown, frontoclypeus glossy dark
brown, genae yellowish. Cervical sclerites, pronotum,
mesonotum tegula dull dark brown. Antenna brown,
but scape light brown. Palps brown, maxillary palps
3.0 mm; proboscis 2.0 mm, as long as first 3% seg-
ments of maxillaty palp. Forewing 16 mm, uniformly
dark brown, with small hyaline spots at nygma and
m-cu, and large translucent spot at arculus. Front and
middle leg with coxa, trochanter and basal % of femur
dark brown, and distal portion of leg light brown;
hind leg light brown. Modified spur (fig. 26) long
and truncate, about 3x longer than adjacent spur,
with apex more than 2x as wide as base, and minute
apical points curved mesad. Genitalia (fig. 26):
Tergum IX short and slightly overhanging segment X
in lateral view, posterior margin rectangular with lat-
eral edges curved in dorsal view; sternum IX with el-
ongate mesosuperior process, extending above % of
phallus in lateral view. Segment X acuminate with
blunt apex in lateral view; broad ellipsoidal with apex
slightly incised in dorsal view. Preanal appendages tri-
angular with corners rounded and dorsal margin
slightly concave in lateral view, ventroposterior mar-
gin curved dorsad, apex broad and blunt. Inferior ap-
pendages capitate with curved truncate apex in lateral
view.
Distribution. — Malaysia: Sabah.
Remarks. — This species was previously only
known from the unique type from Sabah. Two addi-
tional male specimens were found in the material bor-
rowed from BPBM and BMNH. The male is distin-
guished by having modified spur long and truncate,
about 3x longer than adjacent spur, with apex more
than 2X wider than base, and with minute apical
points curving mesad.
Dipseudopsis flavisignata (McLachlan, 1866)
(fig. 27)
Nesopsyche flavisignata McLachlan, 1866: 269, pl. 17: fig. 6,
pl: 19 fig. 6, Type d*: INDONESIA: Sulawesi: Ujung
Pandang, ‘Celebes Macassar’, Wallace coll. (BMNH). The
type was mounted on two separate slides by Mosely
116
(1933), one slide with wings mounted dry, and the sec-
ond slide with body mounted in balsam, but prothorax
absent.
Dipseudopsis flavisignata (McLachlan, 1866). — Mosely
1933: 499-502, figs. 1-7. Fischer 1962: 9. Fischer 1972:
4.
Male. — Head dorsum dark brown, yellowish
brown between antenna, frontoclypeus dark brown.
Antenna base of scape yellowish brown and remain-
der dark brown. Maxillary palps 2.3 mm dark brown,
labial palps 0.8 mm yellowish. Proboscis 1.7 mm
about as long as first four segments of maxillary palp.
Mesothorax and abdomen black. Forewing 12 mm,
similar to that of D. spectabilis (fig. 9), dark brown
background with large yellowish spots: 1) irregular
band on anterior margin tapering toward base, 2)
subapical C-shaped spot having distal side open and
filled with dark narrow stripe along M, (but in de-
scription by Mosely (1933: fig. 1) the latter is depict-
ed as an irregular O-shaped spot), 3) semicircular ir-
regular band through anal region, running from base,
distad through anal loops and thyridial cell to arculus.
Front leg with coxa, trochanter and basal portion of
femur dark, apical portion yellowish; Middle and
hind leg with similar colour pattern but lighter.
Modified spur (fig. 27) nearly as long as first tarsal
segment, basal 4 unmodified, distal slender twisted
blade, bearing a small tooth at base, a ridge at mid-
length and apical % flattened like a blade. Genitalia
(fig. 27): Tergum IX posterior margin broad at base
in dorsal view, but abruptly tapering into slender fin-
gerlike extension, apical 4 overhanging segment X in
lateral view. Segment X long ovoid projection taper-
ing slightly posteriad in dorsal view. Preanal appenda-
ges with dorsal margin nearly horizontal, meeting
curved ventroposterior margin at small dorsoapical
lobe in lateral view. Inferior appendages similar as in
D. spectabilis, heavily sclerotized, and darker brown
than other appendages, basal 4 angled dorsally, and
apical % fingerlike and angled more posteriorly in lat-
eral view.
Distribution. — Indonesia: Sulawesi.
Remarks. — This species is only known from the
unique type. It is a sibling species to D. spectabilis, but
differs by having the forewing with a yellowish C-
shaped subapical spot, almost closed except for a nar-
row dark strip along M,, and by having the modified
spur with one short subapical point distant from api-
cal point.
Dipseudopsis flinti sp. n.
(figs. 11, 28)
Type material. — Holotype d : MALAYSIA: West Malaysia:
WEAVER & MALICKy: Dipseudopsis from Asia
| 281
29E
29C
Figures 26-29. — Dipseudopsis, A-G, male genitalia: A, lateral; B, preanal appendages, segments IX and X, dorsal; C, inferior
appendages, ventral; D, phallus, lateral; E, phallus, ventral; F, segments IX, X, dorsal; G, ventral. I-J, male modified apical
spurs of hind tibia, ventral. Fig. 26, D. elongata Banks. Fig. 27, D. flavisignata (McLachlan). Fig. 28, D. flinti, n. sp. Fig. 29,
D. immaculata Ulmer.
1177
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Selangor, Kuala Lumpur, 21.1V.1931, H. M. Pendlebury, (-
MCZC 33382).
Male. — Head completely glossy dark brown, al-
most black, antennae and mouthparts dark brown.
Maxillary palps 2.3 mm, proboscis 1.3 mm, as long as
first three segments of maxillary palp. Pronotum dull
dark brown, except anterior margin light, mesono-
tum and tegula dark brown and metanotum light
brown. Forewing 12 mm, dark brown except for slen-
der light brown stripe along posterior margin, run-
ning from cell 3A and expanded at arculus; cells be-
low M, and Cu, light brown distally. Front leg with
coxa, trochanter and basal % of femur dark brown,
distal portions of leg light brown; middle leg with
coxa dark brown and distal segments light brown;
hind leg light brown. Modified spur (fig. 28) about
2x longer than adjacent spur, having less than apical %
bifid with two short apical points parallel and angled
distad toward tarsus. Genitalia (fig. 28): Tergum IX
posterior margin broadly bilobed in dorsal view; IX
sternum base rectangular in lateral view, mesosuper-
ior process with blunt dorsal lobe and slender posteri-
or projection with apex curved anteriad. Preanal ap-
pendages trapezoidal with dorsal margin slightly
inclined dorsad toward blunt distal apex in lateral
view, ventral margin curved gradually into posterior
margin. Segment X about % as wide as tergum IX in
dorsal view, somewhat rectangular with posterior
margin broadly bilobed. Inferior appendages capitate
with apical % ellipsoidal and about 3x as thick as base
in lateral view, lateral arms slender and fingerlike with
truncate apex in ventral view.
Etymology. — Named after Oliver S. Flint, Jr.
Distribution. — Malaysia: West Malaysia.
Remarks. — This species is only known from the
unique type. The modified spur of the male is similar
to that of D. stabatensis, but it differs by having two
apical points parallel and angled toward the tarsus,
the adjacent spur is relatively shorter, about % as long
as the modified spur; in D. stabatensis the apical
points of the modified spur are not parallel but are di-
rected in opposite directions, and the adjacent spur is
relatively longer, % as long as the modified spur. Also,
D. flinti differs by having the inferior appendage cap-
itate in lateral view, with the apical % ellipsoidal and
about 3x as broad as the base.
Dipseudopsis immaculata Ulmer, 1905
(fig. 29)
Dipseudopsis immaculata Ulmer, 1905: 92-93, 96, figs. 68b,
69, Type d, Borneo’ in alcohol (MNHN). — Fischer 1962:
118
9; Weidner 1964: 73; Fischer 1972: 4; Malicky &
Weaver 1988: 4, fig. la-e.
Dipseudopsis moesta Banks, 1931b: 400, 401, fig. 9, Type
d*: MALAYSIA: West Malaysia: Johor, Lubok,
Kedondong, NW Gunung Ledang (Mt Ophir), Lalang &
Belukar, H. C. Abraham (MCZC 16481). — Fischer 1962:
11. Syn. n.
nec Dipseudopsis moesta. — Ulmer 1951: 128 [reidentifica-
tion of Ulmer 1905, fig. 75b sub nom. D. stellata) [= D.
benardi].
Specimens examined. — INDONESIA: Sumatra: 2d, Karny,
Pakan Baroe, at light, 23.X.1925 (RMNH). 1d, Huta
Padang, 20.1.1990, Diehl (CLHM). 1¢, Dolok Merangir,
15.V1.82-4.1.1983, Diehl (CLHM). MALAYSIA: Sarawak: 16,
Merirai Valley near Kapft, 180m, 28.VII.-6.VIII.1958, T.
C. Maa (BPBM). West Malaysia: 1d 19, Pahang, Rompin
Mining Co. railway track, 3lkm, Petoh Swamp,
11.XII.1960 (BPBM).
Male. — Head dorsum, cervical sclerite and prono-
tum light orangish brown; frontoclypeus mostly dark
brown, but lateral margins yellowish brown; genae
yellowish brown except dark brown ventral spike.
Antennae light brown. Mouthparts brown, maxillary
palps 2.2 mm; proboscis 1.2 mm, about as long as
first three segments of maxillary palps. Mesonotum
and tegula dark brown, about as dark as eyes. Legs
light brown. Modified spur (fig. 29) slightly longer
than adjacent spur, apical % modified into twisted
point with setose basal pocket; apex almost truncate,
but lateral side curved and other side with short point
directed toward tarsus. Tegula dark brown. Forewing
11-12 mm, almost uniformly brown, except for hya-
line spots at m-cu and arculus. Genitalia (fig. 29):
Tergum IX triangular in dorsal view, with about % of
posterior overhanging segment X in lateral view.
Segment X thumblike in lateral view, extending pos-
teriad as far as inferior appendage; broadly cordate in
dorsal view with apex slightly incised. Preanal appen-
dages rhomboidal or sometimes simply rounded in
lateral view. Inferior appendages oblong and truncate
in lateral view, ellipsoidal in ventral view, but api-
comesal margin incised with blunt subapicomesal
tooth.
Distribution. — Indonesia: Sumatra; Malaysia:
Sarawak, West Malaysia.
Remarks. — The male of this species has a modified
spur which is similar to that of D. doehleri and D.
nebulosa, but can be distinguished from these by its
almost uniformly dark brown forewing. It also differs
by having the male genitalia with more than % of ter-
gum IX overhanging segment X, and the inferior ap-
pendage with a blunt subapicomesal tooth in ventral
view. A lighter phase is noted in male specimens from
Borneo, having apical region of the forewing brown,
remainder nearly translucent except for several dark
WEAVER & MALICKY: Dipseudopsis from Asia
Figures 30-33. — Dipseudopsis, A-G, male genitalia: A, lateral; B, preanal appendages, segments IX and X, dorsal; C, inferior
appendages, ventral; D, phallus, lateral; E, phallus, ventral; F, segments IX, X, dorsal; G, ventral. I-J, male modified spurs ap-
ical spurs of hind tibia, ventral. Fig. 30, D. indica McLachlan. Fig. 31, D. infuscata McLachlan. Fig. 32, D. knappi Schmid
and Denning. Fig. 33, D. lamellata Martynov.
UG)
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
veins. A slight variation also is observed in males from
Sumatra, having forewing with narrow lighter strip
along anterior margin and indistinct translucent area
in outer X of wing, and preanal appendage more no-
ticeably incised distally. Regarding specimen dete-
rmined as D. stellata by Ulmer (1905: fig. 75b) and
later determined as D. moesta by Ulmer (1951), cf.
‘Remarks’ of D. benardi above.
Dipseudopsis indica McLachlan, 1875
(fig. 30)
Dipseudopsis indica McLachlan, 1875: 18-20, pl. 2: figs. 13-
14 (indicus), Type “Habitus in India orientali.’ Lectotype
3: INDIA, designated by Kimmins (1957: 101). — Fischer
1962: 10; Fischer 1972: 4; Higler 1992: 80.
Dipseudopsis buddha Banks, 1913: 239, pl. 23: fig. 8, pl. 24:
fig. 19, Type d*: INDIA: West Bengal, Chapra,
Mackenzie, no date (MCZC 11757). — Fischer 1962: 5;
Higler 1992: 80. Syn. n.
Dipseudopsis indica sindica Martynov, 1935: 159, 206, fig.
59a-b, Type: Pakistan: ‘d. Bubak, Sind, from pools.
12.X1.1927. B. Prashad and B. N. Chopra.’ — Fischer
1962: 10; Fischer 1972: 4. Syn. n.
Specimens examined. — BANGLADESH: 18d 369,
Chicknagul, 10mi N Sylhet, 10m, 30.IX.61, Ross &
Cavagnaro (CASC). 5d 69, Cormilla, 50ft, 23.IX.61, Ross
& Cavagnaro (CASC). INDIA: Bihar: 16, Pusa, at light, 22.-
VIII.08 (MCZC). 18, Pusa, IV.1919, M. A. Husain (BMNH).
1d, Pusa, 2.VIII.1924, Mukerjee (BMNH). Orissa: 2, det.
Martynoy, Balighai, near Puri, 16-20.VIII.11 (ZRAS). 16,
Cuttack, 6.III.1944, at light (BMNH). 2d, Bhubaneswar,
11.1983, J. Olah (CLHM). West Bengal: 1d, Chapra,
Mackenzie (MCZC). 66 322, Malda 110m, 28.X.61, Ross
& Cavagnaro (CASC). 2d 29, 10mi SE Asansol 175m,
6.XI.61, Ross & Cavagnaro (CASC). Pakistan: Sind: 16,
Makli, near Thatta, 22.IX.1976, Hevel & Dietz (USNM). 24
19, Makli, near Hyderabad, 24.IX.1976, Hevel & Dietz (-
USNM).
Male. — Colour dull brown, with eyes dark brown.
Pronotum lighter brown than head and mesonotum.
Maxillary palps 2.0 mm; proboscis 0.8 mm, as long as
first 34 segments of maxillary palp. Legs light brown.
Modified spur (fig. 30) with apical 4 modified, subdi-
vided into two acuminate branches, first branch bifid
with apical points unequal and both directed distally,
second branch curved, directed laterally with minute
point at its base. Forewing 11-14 mm, mostly uni-
form dull brown, but costal margin lighter brown,
with arc of faint translucent spots distad of cord, cen-
tral spot at m-cu, and two translucent spots at arcu-
lus. Genitalia (fig. 30): Tergum IX only slightly
overhanging segment X in lateral view, posterior mar-
gin broadly curved in dorsal view, but middle slightly
protruding; sternum IX short, with mesosuperior
process acuminate in lateral view, extending above
basal % of phallus. Segment X dorsal margin inclined
ventrad with blunt apex lobiform in lateral view, el-
120
lipsoidal and long in dorsal view. Preanal appendages
with dorsal margin straight and horizontal in lateral
view, ventroposterior margin curved apicodorsal an-
gle squarish. Inferior appendages capitate in lateral
view, with short ventrobasal shelf, narrow middle sec-
tion and broad squarish apex; lateral arms with api-
comesal tooth in ventral view.
Distribution. — Bangladesh; India: Bihar, Orissa,
West Bengal; Pakistan: Sind.
Remarks. — This species is easily recognized by the
complex male modified spur, having two branches,
one branch directed distally and bifid, and the other
branch curved laterally and acuminate, but with a
minute point at its base. Examination of the type of
D. buddha revealed that it is identical to D. indica.
Based on its original description, the subspecies D. i.
sindica is recognized as a junior synonym of D. indi-
ca.
Dipseudopsis infuscata McLachlan, 1875
(fig. 31)
Dipseudopsis infuscata McLachlan, 1875: 15-16, pl. 2: figs.
6-10, Holotype d*: INDONESIA: Sulawesi, Ujung
Pandang ‘Piepers Celebes Macassar’ (RMNH). — Fischer
1962: 10; Fischer 1972: 4; Malicky & Weaver 1988: 3,
fig.4a-e.
nec Dipseudopsis infuscata. — Ulmer 1905: 96, fig. 75d;
Ulmer 1951: 128; Malicky 1984: 214, 216
Specimens examined. — INDONESIA: Java, Jakarta ‘Bata-
via: 1d, 11.1908; 1d, VIII.1907; 18, IX.1907 (RMNH);
14, 1.1908, Jacobson (ZMAN). Kalimantan: 16 39, Telang
X.1881 (MCZC). 1d, West Java, [Piepers?], McL. coll
(BMNH). Sumatra: 2d, Palembang, 13.V.1972, Diehl
(CLHM).
Male. — Head and antennae brown. Maxillary
palps 1.6-1.8 mm, proboscis 0.5-1.0 mm as long as
first two or three segments of maxillary palp. Antenna
yellowish brown with fine brown setae that become
less dense apically so that distal part appears more yel-
lowish. Forewing 12 mm, brown with small translu-
cent spot distal to r, hyaline spots at m-cu and arcu-
lus. Modified spur (fig. 31) slightly longer than
adjacent spur, bifid with two acuminate apical points,
longer point straight directed distally and almost % as
long as long as spur, shorter point curved with apex
directed mesad. Genitalia (fig. 31): Tergum IX with
posterior margin rounded in dorsal view, with slight
protrusion in middle, short and not extended over
segment X in lateral view. Segment X with basal # of
dorsal margin inclined ventrad apical portion direct-
ed posteriad, having apex truncate, in lateral view;
dorsal and ventral margins narrowing toward apex,
with apex truncate and slightly emarginate in dorsal
view. Preanal appendages with posterior margin in-
cised in lateral view. Inferior appendages with dorsal
and ventral margins nearly parallel and curved dorsad
in lateral view, having thumblike apex; base with me-
sal margin squarish in ventral view, distal portion
long with mesal margin slightly convex, apicolateral
margin rounded, and with apicomesal point.
Distribution. — Indonesia: Java, Kalimantan,
Sulawesi, Sumatra.
Remarks. — This species is similar to D. robustior,
but differs by having the two apical points of the
modified spur with the longer point straight and
slightly shorter than the unmodified base of the spur,
and the shorter point curved with its apex directed
mesad. All specimens determined as D. infuscata by
Malicky (1984) from South Andaman Island are now
considered to be D. robustior andamanensis Weaver &
Malicky, n. ssp. The specimen referred to by Ulmer
(1905: fig, 75d, 1951) is D. contorta, not D. infuscata
(cf. D. contorta above).
Dipseudopsis knappi Schmid & Denning, 1979
(fig. 32)
Dipseudopsis knappi Schmid & Denning, 1979: 246, 248,
fig. Sa-e, Holotype d: THAILAND: Tha Phra, Khom Kaen,
17.V.1968, blacklight trap, F. W. Knapp (CASC).
Male. — Body brown. Forewing 13 mm, brown
with scattered translucent spots. Modified spur (fig.
32) trifid with long curved acuminate point and
slightly shorter fingerlike lobe bearing minute subap-
ical point. Genitalia (fig. 32): Tergum IX broadly
rounded in dorsal view, short in lateral view; sternum
IX with mesosuperior process triangular in lateral
view. Segment X slender with apex distinctly bilobed
in dorsal view, lateral margins sinuate, base of dorsal
margin slightly inclined dorsally distad in lateral view,
apical 4 horizontal with ventral margin concave, apex
slender and fingerlike. Preanal appendages trapezoid-
al with corners rounded and dorsal margin inclined
dorsad in lateral view, having blunt apicodorsal lobe.
Inferior appendages slightly clavate with basodorsal
shelf in lateral view, middle portion constricted, and
having clavate apex.
Distribution. — Thailand.
Remarks. — This species is only known from the
type series from northeastern Thailand. The modified
spur of D. knappi is trifid and somewhat similar to
that of D. spectabilis, but differs by having second
longest process of the spur lobiform, rather than
pointed. The male genitalia differ by having segment
WEAVER & MALICKY: Dipseudopsis from Asia
X with a bilobed apex. The diagnosis of the male giv-
en above is based on the original species description
(Schmid & Denning 1979).
Dipseudopsis lamellata Martynov, 1935
(fig. 33)
Dipseudopsis lamellata Martynov, 1935: 160-161, figs. 60a-
b, 61, Type series: BURMA: Myitkyina Dist: 16 129,
Lonton, W shores Indawgyi Lake, U. Burma, 18-
31.X.1926, B. N. Chopra; Syntype 19 *: Kamaing, U.
Burma, X.1926, B. N. Chopra (NZSI); INDIA: Assam:
Mangaldai, 26.26N 92.02E, 16-18.X.1910, S. W. Kemp.
Fischer 1962: 10; Higler 1992: 80.
Specimens examined. — BANGLADESH: 1d, Chicknagul,
10mi N Sylhet, 10m, 30.IX.61, Ross & Cavagnaro (CASC).
23 49, Harbang For. 6mi N Chiringa, 50ft, 18.IX.61,
Ross & Cavagnaro (CASC). 1d, Umgebung Dacca, 1-
15.V.1976, Dietz (CLHM). INDIA: Assam: 26 32, Kohara,
Kaziranga, 110m, 26.36N 93.28E, 7-16.X.61, Ross &
Cavagnaro (CASC). Orissa: 28, Bhubaneswar, 11.1983, J.
Olah (CLHM).
Male. — Colour brown. Forewing 13-15 mm,
colouration similar as in D. indica, mostly brown, but
with arc of five faint translucent spots distad of cord,
central hyaline spot at m-cu, and broad translucent
spot at arculus. Modified spur (fig. 33) 2X longer than
adjacent spur, bearing two distal processes, longer
process slender twisted and fingerlike, shorter process
slender acuminate and bent laterad at right angle.
Genitalia (fig. 33): Tergum IX broad triangular in
dorsal view, apex slightly overhanging segment X in
lateral view; sternum IX short, with mesosuperior
process having blunt dorsal lobe and slender ventr-
oposterior process in lateral view. Segment X dorsal
margin concave, ventral margin convex, apical part
narrow and curved dorsad in lateral view; lateral mar-
gins slightly convergent at basal %, distal portion elon-
gate ellipsoidal, with apex slightly incised in dorsal
view. Preanal appendages slender and somewhat trap-
ezoidal in lateral view, with posterior margin strongly
incised, bearing two blunt apical points. Inferior ap-
pendages capitate in lateral view, with base constrict-
ed, having short slender basal projection and widen-
ing into rhomboid apex; basomesal margin with deep
curved incision in ventral view, distal portion wid-
ened, having apex with lateral margin curved and
with short apicomesal point.
Distribution. — Bangladesh; Burma; India: Assam,
Orissa.
Remarks. — This species differs from other species
of Dipseudopsis by having a unique modified spur, bi-
fid, with a long slender fingerlike twisted process, and
with an acuminate process angled laterad.
121
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figures 34-37. — Dipseudopsis, A-G, male genitalia: A, lateral; B, preanal appendages, segments IX and X, dorsal; C, inferior
appendages, ventral; D, phallus, lateral; E, phallus, ventral; F, segments IX, X, dorsal; G, ventral. I-J, male modified spurs ap-
ical spurs of hind tibia, ventral. Fig. 34, D. lucasi, n. sp. Fig. 35, D. maculata Ulmer. Fig. 36, D. malaisei, n. sp. Fig. 37, D.
martynovi, n. sp.
122
Dipseudopsis lucasi sp. n.
(fig. 34)
Type material. — Holotype d: INDIA: Karnataka:
Shimoga Dist., Agumbe Ghat, 2000ft, 1974, Lucas (CLHM).
Paratypes: 76 39, Someshwar, 27.1.1959, F. Schmid
(CNCI). 16, Nagodi, 28.1.1959, F. Schmid (cncı). 1d,
Kerala: Sangalipalam, 20-21.XII.1961, F. Schmid (cncı).
Male. — Head brown, eyes darker brown, fronto-
clypeus brown, antennae brown but ventral sides
lighter brown, palps brown, maxillary palps 2.8 mm,
labial palps 1.0 mm, proboscis 1.2 mm, almost as
long as first three segments of maxillary palp.
Pronotum dark brown, but lateral sides lighter, mes-
onotum and tegula dark brown. Forewing 14-17 mm
unicoloured dark brown except small hyaline spots at
m-cu and arculus. Legs with coxae dark brown, distal
segments light brown. Modified spur (fig. 34) slight-
ly longer than adjacent spur, apical X modified with
short curved subapical shelf, and long slender acumi-
nate curved point directed distad. Genitalia (fig. 34):
Tergum IX with posterior 4 overhanging X in lateral
view, posterior margin broadly rounded in dorsal
view; sternum IX mesosuperior process with middle
constricted and apex curved abruptly dorsad in lateral
view. Segment X thumblike and directly posteriad in
lateral view; elongate ovoid, with apex slightly incised
in dorsal view. Preanal appendages short, extending
to middle of segment X in lateral view, posterior mar-
gin broadly rounded, almost semicircular but with
blunt apicodorsal angle. Inferior appendages basal %
thick and broadened toward middle in lateral view,
apical 4 tapered to truncate apex; lateral margin
rounded and mesal margin almost straight in ventral
view, but with obtuse apicomesal angle and minute
basomesal angle.
Etymology. — Named after the collector J.A.W.
Lucas.
Distribution. — India: Karnataka, Kerala.
Remarks. — The male genitalia of D. lucasi are sim-
ilar to those of D. doehleriand D. nebulosa, having ab-
breviated preanal appendages. However, D. lucasi dif-
fers by having forewing almost unicoloured dark
brown, and lacking the contrasting pattern, as in the
forewings of D. doehleri and D. nebulosa (figs. 5, 7).
Dipseudopsis maculata Ulmer, 1907
(fig. 35)
Dipseudopsis maculata Ulmer, 1907a: 37-39, fig. 55a-b,
Type d*: MALAYSIA: Sabah: Sandakan (BMNH). Fischer
1962: 11. Fischer 1972: 5.
WEAVER & MALICKY: Dipseudopsis from Asia
Specimens examined. — MALAYSIA: Sabah: 1d, Tawau,
Quoin Hill, Cocoa Res. Sta., 1.X.1962, Y. Hirashima
(BPBM). 16, Tawau, Quoin Hill, 15-20.VII.1962, Y.
Hirashima (BPBM). 1d, Sepilok For. Res., Sandakan Bay, 1-
10m, 27.X.1957, J. L. Gressitt (BPBM).
Male. — Head dorsum with anterior % glossy dark
brown, almost black, and posterior warts light brown,
frontoclypeus glossy dark brown; antennal scape and
pedicel light brown, flagellum darker, palps dark
brown, maxillary palps 3.6 mm, proboscis 2.2 mm, as
long as first four segments of maxillary palp.
Pronotum, mesonotum and tegula dark brown, but
metathorax and abdomen yellowish brown. Forewing
17-18 mm, veins dark brown against brown and
translucent background, large translucent triangular
spot between R, and Cu,, also translucent spots at ar-
culus, discal cell, and area distal to cord, and dark
band distal to hyaline spot at m-cu. Fore leg coxa to
basal 4 of femur dark brown, distal % of femur and
other distal leg segments yellowish brown; mid leg
coxa dark brown, distal leg segments yellowish
brown; hind leg yellowish brown. Modified spur (fig.
35) 2x longer than adjacent spur, apical 4 modified,
into two unequal short nearly parallel points curved
laterally away from tarsus. Genitalia (fig. 35):
Tergum IX distal 4 overhanging segment X and with
broad apex in lateral view, posterior margin broadly
rounded and bilobed in dorsal view. Sternum IX me-
sosuperior process triangular with blunt dorsal angle
and acute posterior angle in lateral view. Segment X
ventral margin straight and horizontal, dorsal margin
slightly concave and inclined ventrad, apex rounded;
ellipsoidal about 4 as wide as tergum IX in dorsal
view. Preanal appendages broad and triangular with
corners rounded in lateral view, dorsal and ventropo-
sterior margins slightly irregular, apicodorsal apex
blunt. Inferior appendages inclined dorsad and wid-
ening apically, having apex truncate with rounded
corners in lateral view, dorsal margin straight and
longer than ventral margin; ventral margin with basal
% having protruding shelf extended as short free lobe,
oblique angle at midlength, and distal % inclined dor-
sad; distal process thumblike in ventral view, mesal
margin with deep narrow curved basolateral notch,
and apex rounded.
Distribution. — Malaysia: Sabah.
Remarks. — The male of this species has a unique
modified spur with a prominent apical point extend-
ing laterally from a blunt apical shelf, and a shorter
subapical point curved laterally, having both points
curved distad. D. maculata and D. nieuwenhuisi ap-
pear to be closely related based on similar colouration
in the male forewing (cf. fig. 6 of D. nieuwenhuisi),
and similar characteristics in the male genitalia. D.
123
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
maculata differs by being smaller in size, and by hav-
ing male modified spur with only two apical points,
rather than having three as in D. nieuwenhuisi.
Dipseudopsis malaisei sp. n.
(fig. 36)
Type material. — Holotype d : BURMA: Washaung, 600ft,
13.VII.1934, Malaise (NHRS). Paratypes: 34, same data,
Malaise (NHRS).
Male. — Head dorsum and frontoclypeus dark
brown, nearly black, posterior warts brown.
Antennae, mouthparts, legs and sclerotized parts of
thorax unicoloured brown. Maxillary palps 2.2-2.5
mm; proboscis 1.2-1.8 mm, as long as first four seg-
ments of maxillary palps. Forewing length 13-15
mm, dark brown except for small hyaline comma at
m-cu. Modified spur (fig. 36) slender, slightly curved
and longer than adjacent spur, with two small spiral-
ling apical points. Abdomen yellowish brown.
Genitalia (fig. 36): Tergum IX broadly rounded in
dorsal view, with distal 4 overhanging segment X in
lateral view; sternum IX with mesosuperior process
fingerlike and % as long as segment X in lateral view.
Segment X long and fingerlike, tapering slightly dis-
tad and apex incised in dorsal view. Preanal appenda-
ges oblong, with posterodorsal margin ellipsoidal in
lateral view. Inferior appendages curved dorsad and
broad in lateral view; broad with lateral and mesal
margins parallel, and apices slightly bent mesad in
ventral view.
Etymology. — Named after the collector R. Malaise.
Distribution. — Burma.
Remarks. — This species is only known from the
type series. The male has segment X long and slender
and modified spur with two small apical spiralling
points similar as in D. benardi. However, D. malaisei
differs by having male with preanal appendages ob-
long and ellipsoidal in lateral view, inferior appenda-
ges broad and flat, and dark brown forewings without
large translucent spots, similar to D. benardi (fig. 10).
Dipseudopsis martynovi sp. n.
(fig. 37)
Type material. — Holotype d: PHILIPPINES: Sulu:
Tarawakan NE, 12.11.1957 Yoshio Kondo (BPBM).
Paratypes: Tawitawi, Tarawakan, north of Batu Batu,
Noona Dan Exp. (ZMUC): 106, 21.X.1961; 4d,
20.X.1961; 24, 13.X1.1961; 24 : 14.X1.1961.
Male. — Head dark brown, but posterior warts light
124
brown; frontoclypeus with ventral % light brown, dor-
sal 4 dark brown; antennae light brown, palps light
brown, maxillary palps 2.3 mm, proboscis 1.2 mm, as
long as first three segments of maxillary palp.
Pronotum light brown. Mesonotum and tegula dark
brown. Forewing 13 mm, dark brown with C and
small light brown stigma and small hyaline spots at
m-cu and arculus. Legs with coxae dark brown, distal
segments light brown. Modified spur (fig. 37) short
and stout, about 4x, as long as basal width apical %
modified with three broad points clustered together.
Genitalia (fig. 37): Tergum IX posterior 4 extended
over segment X in lateral view, broad and triangular
in dorsal view; sternum IX mesosuperior process
straight rodlike with apex curved dorsad in lateral
view. Segment X dorsal margin slightly concave and
inclined ventrad in lateral view, ventral margin
curved ventrad at middle and curved dorsad distad,
apex blunt; ellipsoidal and about % as wide as tergum
IX in dorsal view. Preanal appendages slightly shorter
than segment X in lateral view, dorsal margin slightly
sinuate at base, with broad dorsoapical curve, ventral
margin nearly straight inclined dorsad, posterior mar-
gin with shallow incision. Inferior appendages finger-
like and inclined dorsad in lateral view, with narrow
basoventral shelf, squarish with distal arms parallel
and straight in ventral view, lateral apex rounded, ba-
somesal margin with square-angled basal shelf, and
lateral notch and apicomesal angle both square.
Etymology. — Named after Andrei Vasilievich
Martynov.
Distribution. — Philippines: Sulu, Tawitawi.
Remarks. — This species is superficially similar to
D. recta and D. triclavata, having modified spur tri-
fid. However, it differs by having the spur short and
stout, about 4X longer than basal width, with apical %
of spur modified, and inferior appendage fingerlike in
lateral view.
Dipseudopsis modesta Banks, 1911
(fig. 38)
Dipseudopsis modesta Banks, 1911: 105-106, pl. 6: fig. 2,
Type 4 *: INDIA: Bihar: Pusa, 21.IX.08, T. N. J. (MCZC
11756). — Fischer 1962: 11; Fischer 1972: 5; Higler
1992: 80.
Dipseudopsis pallida Martynov, 1935: 166-167, fig. 70a-b,
Type series: 1d 1 9 in copula, Pusa, 31.111.1911-4.IV, F.
H. Gravely (probably NZSI). — Fischer 1962: 14; Higler
1992: 80. Syn. n.
Specimens examined. — INDIA: Assam: 38 39, Kohara,
Kaziranga, 110m, 26.36N 93.28E, 16.X.61, Ross &
Cavagnaro (CASC). 1d 39, 13mi SW Kochugaon, 50m,
17.X.61, Ross & Cavagnaro (CASC). 2d 29, Meleng For.,
Mariani, 110m, 26.39N 94.18E, 10.X.61, Ross &
Cavagnaro (CASC). 2d 39, Garampani Res. 10mi S
Gologhat, 110m, 9.X.61, Ross & Cavagnaro (CASC). Assam:
16 22, ‘Doom Dooma Dum Duma, 27.34N 95.34E,
VI.1943, E. Hardy (USNM). ‘U.D.M.N.C.H. [United District
of Mikir and North Cachae Hills]: 16, Kalanga, 1.V.1960,
F. Schmid (encı). Bihar: 4d 29, Pusa (MCZC). 2d, Pusa
(BMNH). Orissa: 2d, Bhubaneswar, 21.11.1985, J. Olah
(CLHM). West Bengal: 34, Chapra, Mackenzie (MCZC).
Male. — Head glossy dark brown; frontoclypeus, an-
tennae and palps dark brown; maxillary palps 2.6 mm,
proboscis 1.0 mm, almost as long as first three seg-
ments of maxillary palp. Pronotum, mesonotum and
tegula dark brown. Forewing 12-15 mm, dark brown,
except for hyaline lunula at m-cu and small translu-
cent spot at arculus. Legs light brown slightly darker
basad. Modified spur (fig. 38) slightly shorter than ad-
jacent spur with two minute apical points (cf.
‘Remarks’ below). Genitalia (fig. 38): Tergum IX tri-
angular with acute apex slightly inclined dorsad and
distal 4 overhanging segment X in lateral view; poste-
rior margin broadly rounded in dorsal view; sternum
IX mesosuperior process with short broad dorsal trian-
gle, posterior process narrow having dorsal margin ir-
regular and slightly concave, and ventral margin
curved dorsad in lateral view, extending above basal
of phallus. Segment X long slender and apex bilobed
with deep mesal notch in dorsal view; fingerlike and
slightly tapered in middle, extending posteriad distal
to inferior appendages in lateral view. Preanal appen-
dages trapezoidal, posterior margin slightly incised in
lateral view. Inferior appendages slender and long in
lateral view, extending dorsad toward apex of preanal
appendage, dorsal margin curved and concave, ventral
margin irregular; lateral margins curved and apicome-
sal margin straight with apical point in ventral view,
mesal margin with curved notch in basolateral corner.
Distribution. — India: Assam, Bihar, Orissa, West
Bengal.
Remarks. — This species can be distinguished by its
unique male modified spur, having a short inconspic-
uous apical point directed distally and a long slender
point curving halfway around the apex; the straight
point is minute and lightly sclerotized, and without
clear magnification it appears to be a small lobe. This
species is similar to D. benardi, but it differs by hav-
ing male modified spur with smaller apical points and
inferior appendages narrower in lateral view. We have
observed some variation in the shape of the preanal
appendages. Comparison of the type of D. modesta
with the original description of D. pallida, reveals that
these species are conspecific, and thus the latter is rec-
ognized as a junior synonym.
WEAVER & MALICKY: Dipseudopsis from Asia
Dipseudopsis morosa Banks, 1924
(fig. 39)
Dipseudopsis morosa Banks, 1924: 450, pl. 3: fig. 31. —
Fischer 1962: 11; Fischer 1972: 5.
nec Dipseudopsis morosa. — Schmid 1958: 13, 24, 27, 89, pl.
15: figs. 12-13.
Lectotype designation. — d*: PHILIPPINES:
Mindanao: Dapitan, Baker (MCZC 14826). 16*:
Basilan: probably syntype but without type label,
Baker (USNM). The type series: 34 (MCZC) is mixed
and comprises three different species; in addition to
the lectotype, the d * syntype from Luzon is D. colla-
ris, and the d * syntype from ‘Borneo’ Kalimantan is
D. stabatensis.
Specimens examined. — PHILIPPINES: Mindanao: 16,
Lanao, 4.8 km E of Dansalan, 750m, 11.VI.1958, jungle
along stream, H. E. Milliron (BPBM).
Male. — Head brown with black triangular median
stripe, broad along anterior margin and tapering to
middle of posterior margin, frontoclypeus glossy
black and genae brown, antennae and palps brown,
maxillary palps 2.3 mm, proboscis 1.0 mm, almost as
long as first three segments of maxillary palp.
Pronotum black with posterior dorsal margin brown,
mesonotum and tegula almost black. Forewing 9-10
mm, uniformly dark brown, but small hyaline spots
at m-cu and arculus. Legs with coxae of fore and mid
legs dark brown and distal segments yellowish, hind
leg yellowish. Modified spur (fig. 39) 14x longer than
adjacent spur, apex truncate with two apical points,
both points curved and one point nearly in plane per-
pendicular to longitudinal axis of spur, and other
longer point with apex directed more distad.
Genitalia (fig. 39): Tergum IX not extended over base
of segment X in lateral view, slightly inclined dorsad;
sternum IX mesosuperior process short and blunt in
lateral view, with square dorsoanterior angle, posteri-
or lobe inclined dorsad and extended above basal % of
phallus. Segment X dorsal and ventral margins nearly
parallel and inclined ventrad in lateral view, but api-
cal portion of ventral margin horizontal, apex round-
ed; triangular and with rounded posterior apex in
dorsal view. Preanal appendages broad and somewhat
D-shaped with apex extended dorsoposteriad in late-
ral view. Inferior appendages oblong with distinct
subapical ventral tooth in lateral view, distal portion
capitate in ventral view with apex and basomesal
notch curved.
Distribution. — Philippines: Basilan, Mindanao.
Remarks. — D. morosa has been confused with D.
triclavata, due to similarities of the male modified
spurs. However, these species are not closely related
and are distinguished from each other by characteris-
125
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figures 38-41. — Dipseudopsis, A-G, male genitalia: A, lateral; B, preanal appendages, segments IX and X, dorsal; C, inferior
appendages, ventral; D, phallus, lateral; E, phallus, ventral; F, segments IX, X, dorsal; G, ventral. I-J, male modified spurs ap-
ical spurs of hind tibia, ventral. Fig. 38, D. modesta Banks. Fig. 39, D. morosa Banks. Fig. 40, D. nebulosa Albarda. Fig. 41,
D. nervosa Brauer.
126
tics of the male genitalia, D. morosa having the inferi-
or appendage broader and with a distinct blunt ven-
tral tooth in lateral view, and by having modified spur
with only two apical points. The type series of D. mo-
rosa at the MCZC is mixed, comprising three syntypes,
each a different species. Records of D. morosa from
Sri Lanka by Schmid (1958) are actually D. triclava-
La.
Dipseudopsis nebulosa Albarda, 1881
(figs. 7, 12A, 40)
Dipseudopsis nebulosa Albarda, 1881: 19, pl. 5: fig. 4, Type
2, INDONESIA: ‘in insula Sumatra’. — Fischer 1962: 11-
12; Fischer 1972: 6; Malicky & Weaver 1988: 4, fig. 2a-
ie
nec Dipseudopsis nebulosa. — Banks 1914: 178 (Luzon).
Specimens examined. — BURMA: 2d, Tenasserim, Sukli,
75km E of Moulmein, 600m, 27-31.X.1934, Malaise
(BMNH). 1d, Mekane, 20 km W Myawaddy, Amherst Dist,
700ft, 3.XI.1934, Malaise (NHRS). INDONESIA: Sumatra:
16, W Morton (CNCI). 1d, Tandjong Morawa Serdang
(RMNH). 1d, Loeboek Sikaping, 450m (RMNH). 1d,
Pematang Siantar, 1.VI-12.XII.1985, Diehl (CLHM). 126,
Huta Padang, Diehl (CLHM). MALAYSIA: West Malaysia:
Pahang: 34, Kuala Tahan, at light, Pendlebury (BMNH).
16, Sungei Tembeling, 18.XI.1922, Pendlebury (MCZC).
18, Kuala Teku, 500ft, 5.XII.1921, Pendlebury (MCZC).
Selangor: 126 69, Kuala Lumpur, 15-31.X11.1958, Quate
(BPBM). THAILAND: 36, upper Pran River, 13-15.1V.1926
(MCZC 16 , BMNH 2d). 26, det. Mosely as D. doehleri, same
data, 15.1V.1927, Landell (BMNH). 1d, Trang, 26.VI.1924,
Evans (MCZC). 28 19, Trang Prov., Khaophappha
Khaochang, 200-400m, 12-13.1.1964, Samuelson (BPBM).
5d, Krachong For. near Trang, 100m, 2.VII.62, Ross &
Cavagnaro (CASC). 1d, 20 km E of Krabi, 10-20.11.1962,
Friedel (CLHM). 1d, Trang, Banchang, 16.V.1924, at light,
Evans (BMNH).
Male. — Body and wings (fig. 7). Head dorsum, an-
tennae, cervical sclerites and pronotum orangish
brown, and contrasting with dark brown mesono-
tum, tegula and forewings (fig. 12A). Frontoclypeus
and mouthparts brown. Fore and mid legs with coxae
and trochanters brown, femora with basal portions
dark brown and apical portions orangish brown, tib-
iae and tarsi orangish; hind leg dark brown. Forewing
12-14 mm, dark brown with translucent spots: long
triangular spot below R, above Cu, from base of Rs to
hyaline lunula at m-cu, translucent oblong spot distal
to cord with apical margin deeply incised, and small
translucent spot at arculus. Modified spur (fig. 40)
with twisted glabrous apical point, about % as long as
spur. Genitalia (fig. 40) similar to D. schmidi.
Tergum IX with posterior margin triangular with
apex curved and lateral sides slightly depressed in dor-
sal view, apical 4 of dorsum overhanging segment X
in lateral view. Sternum IX extended anteriad, with
mesosuperior process globular in lateral view.
WEAVER & MALICKY: Dipseudopsis from Asia
Segment X short stout and ellipsoidal in lateral view,
dorsal and. ventral margins nearly parallel and hori-
zontal, apex bluntly rounded, broad and cordate in
dorsal view, length and width subequal, with apex
slightly incised. Preanal appendages abbreviated in
length, with posterior margin incised. Inferior appen-
dages broad, somewhat ellipsoidal in lateral view, but
dorsal margin slightly concave.
Distribution. — Burma; Indonesia: Java, Sumatra;
Malaysia: West Malaysia; Thailand.
Remarks. — This species is similar to D. doehleri,
but differs by having a translucent spot of forewing
with faded edges, rather than a well defined oval-
shaped spot at the arculus, and genitalia with sternum
IX, segment X, and preanal appendages slightly
broader in lateral view. The modified spur of D. neb-
ulosa is similar to that of D. immaculata and D. doeh-
leri, with a twisted glabrous apical point, having a se-
tose pocket at its base, but it differs by having a longer
glabrous point, about % as long as spur. We have ac-
cepted the description of the male provided by Ulmer
(1951), who associated it with the female type from
middle Sumatra. We have observed that a male spec-
imen from Luzon was misidentified by Banks (1914)
and actually is D. collaris.
Dipseudopsis nervosa Brauer, 1868
(figs. 3, 4, 41)
Dipseudopsis nervosa Brauer, 1868: 267-268, Type 2: PHILI-
PPINES: (Semper) [cf. ‘Remarks’ below]. — Fischer 1962:
12; Fischer 1972: 6.
Dipseudopsis luctuosa Banks, 1913: 178, pl. 8: fig. 3, Type
&*: Luzon: Los Banos, P. L. Baker (MCZC 11769). —
Ulmer 1930: 380, as syn. — Fischer 1962: 12; Fischer
1972: 6.
Dipseudopsis venosa Navás, 1931: 86, Type 2 ‘Filipinas:
Mindanao, Kolambogan, 11.VI.1914. Col. m.’ [We were
unable to locate the female type of D. venosa for examina-
tion, but are relatively certain that this species is synony-
mous with D. nervosa, because only two species of
Dipseudopsis are known from Mindanao, D. modesta and
D. nervosa, and the size of the female type of D. venosa
only agrees with the female of D. nervosa.) — Fischer
1962: 16. Syn. n.
Specimens examined. — PHILIPPINES: Leyte: 3d, Palo (-
BPBM). 1d, Tacloban (USNM). Luzon: 3¢ 39, Los Baños (-
MCZC). 2d 59, Los Baños (USNM). 44 39, Mt Maquiling
(CNCI, USNM). 1d, det. Ulmer (1905), bez. Bilneao, prov.
Laguna, 1876, Laglaize (MNHN). Mindanao: 1d,
Kolambugan (BMNH). 16 (‘Paratypus’ see discussion above)
Mindanao, 1867 (NHMW). 2d, Agusan Los Arcos (BPBM).
19, Agusan Esperanza (BPBM). 38, Agusan Jabonga
Mayogda (Mczc). 26, Bukidnon, Dalongdong, 40km NW
Maramag, Talakag, Urwaldrand, 800m, 7°53’N 124°40’E (-
ZMHB). 16 29, Butuan (MCZC, USNM). 1d, Cotabato,
Kalaong (BPBM). 2d, Sapamoro, Curnan dist. (ZMUC).
127
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Mindoro: 1d, Laguna de Nanjan Soldanski (CLHM).
Negros: 28 39, L. Balinsasayao (BPBM). 16, Sibulan
(BPBM). 26 19, Victorias (MCZC, USNM). Panay: 16, Pr.
Capiz, Libacao (ZSMC). Sibuyan: 16, Magdiwang-
Tampayan, Ga-ong, Pawala River (CLHM).
Male. — Body and wings (fig. 3). Head dorsum
similar to D. morosa, having dark brown median
acute triangle, tapering posteriad, and lateral sides
yellowish brown; frontoclypeus with dark brown me-
dian stripe tapering dorsad and lateral sides yellowish
brown; antennae and palps brown, maxillary palps
2.8 mm, proboscis 1.7 mm, as long as first 3% seg-
ments of maxillary palp. Cervical sclerites mostly dark
brown, but dorsal part yellowish brown. Pronotum
yellowish brown, median furrow and lateral sides
dark brown; propleuron yellowish brown.
Mesonotum, metanotum and tegula dark brown.
Mesothorax, metathorax and abdomen brown dorsad
and yellowish brown ventrad. Forewing 14-17 mm,
dark brown with three translucent spots: 1) basal
stripe between R, and Cu, 2) triangular spot distal to
cord, and 3) round spot at arculus. Fore leg coxa bi-
coloured, basal 4 dark brown and distal % yellowish
brown. Pleuron bicoloured, cervical sclerites and side
of thorax with lower portion dark brown and upper
part yellowish brown. Modified spur (fig. 41) trifid,
with long apical point spiraling distad, minute apical
point, and subapical lobe. Genitalia (fig. 41): Tergum
IX inclined dorsad in lateral view, minute portion of
apex overhanging segment X, posterior margin
broadly curved in dorsal view; sternum IX mesosupe-
rior process oblong in lateral view, dorsal margin
curved. Segment X dorsal margin slightly concave,
strongly inclined ventrad to blunt apex in lateral view;
base broad and narrowing toward oval-shaped apex in
dorsal view. Preanal appendages irregular triangle,
with dorsal margin irregular and curved in lateral
view, ventroposterior margin steeply inclined dorsad,
dorsoposterior apex squarish. Inferior appendages
fingerlike and curved dorsad in lateral view, lateral
margin curved in ventral view, extended arms finger-
like with middle of mesal margin having small semi-
circular incision.
Female. — Body and wings (fig. 4).
Distribution. — Philippines: Leyte, Luzon,
Mindanao, Mindoro, Negros, Panay, Sibuyan.
Remarks. — This species is similar to D. digitata,
but differs by having unique colouration of the male
head and forewings, and by characteristics of the male
hind spur and genitalia.
Brauer (1868) described this species after a female
from the Philippines, and the species epithet, nervosa,
was probably derived in references to the colouration
128
pattern of the female forewing, having dark veins
against a translucent background. The type which
should be at the NHMW is missing. Instead a male
specimen with a hand-written type label reading
‘Paratypus’ is present, but this specimen was not
mentioned by Brauer and thus cannot be regarded as
a syntype. The red ‘paratype’ label was probably add-
ed to this specimen after the species was described.
This specimen cannot be the type because the type
description was clearly based on a female specimen;
also the forewing length of the type should be 19 mm,
but that of the male ‘paratype’ is only 17 mm.
Therefore, we conclude that the type is missing. We
accept the male association of this species by Ulmer
(1929). Furthermore, we are reasonably sure that the
identity of this species is correct because there are no
other species as large as D. nervosa known from the
Philippines., i.e. the other four Philippine species, D.
collaris, D. digitata, D. martynovi, and D. morosa, are
much smaller.
Dipseudopsis nervosella Ulmer, 1905 stat. n.
(fig. 42)
Dipseudopsis sp. — Ulmer, 1905: 96, fig. 75e.
Dipseudopsis morosa. — Ulmer 1951: 128 [misdet.].
Dipseudopsis morosa nervosella Ulmer, 1951: 141-143, pl. 7:
figs. 140-142, Type d* ‘Borneo’ 1886, F. Baczes
(NHMW). — Fischer 1972: 5.
Specimens examined. — MALAYSIA: Sarawak: 16 , at foot
of Mt Dulit, junction of rivers Tinjar & Lejok, 19.VI-
11.1932 (BMNH). 1 2, same data, 30.VIII.1932 (BMNH). 1d,
R. Kapah trib. of R. Tinjar, 25.X.1932, Oxford Univ. Exp.
(BMNH).
Male. — Dorsum of head with curved M-shaped
pattern marked by division between glossy dark
brown, almost black, anterior triangle pointed poste-
riad and dark brown lateral margins, and yellowish
brown area, filling in two triangles of ‘M° and with
acute angle of each triangle pointed toward base of
antenna. Frontoclypeus glossy, almost black, ante-
nnae yellowish brown, cervical sclerites dark brown.
Maxillary palps 3.0 mm, dark brown; proboscis
brown, 1.8 mm, almost a long as first four segments
of maxillary palp; labial palps 1.2 mm. Pronotum
dark brown with slender yellowish brown posterior
edges along median fissure and lateral margins, pro-
pleuron light brown; mesonotum almost black, mes-
opleuron dark brown except epimeron light brown;
metathorax and abdomen yellowish brown. Forewing
15-16 mm, dark brown with golden hairs, and three
large translucent spots, similar to D. nervosa and D.
nebulosa (cf. figs. 3, 7): 1) long band between R, and
Cu, 2) equilateral triangle at arculus, and 3) subapical
area distal to cord with six translucent gaps between
WEAVER & MALICKy: Dipseudopsis from Asia
Figures 42-45. — Dipseudopsis, A-G, male genitalia: A, lateral; B, preanal appendages, segments IX and X, dorsal; €, inferior
appendages, ventral; D, phallus, lateral; E, phallus, ventral; F, segments IX, X, dorsal; G, ventral. I-J, male modified spurs ap-
ical spurs of hind tibia, ventral. Fig. 42, D. nervosella Ulmer. Fig. 43, D. nieuwenhuisi Ulmer. Fig. 44, D. notata (Fabricius).
Fig. 45, D. recta Martynov.
129
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
dark apical veins, and in discal cell. Fore leg and mid-
dle leg with coxae, trochanter and basal portion of fe-
mur dark brown, distal portion of leg yellowish; hind
leg yellowish. Modified spur (fig. 42) 14x longer than
adjacent spur, apex truncate with two apical points,
both points curved in plane perpendicular to longitu-
dinal axis of spur (cf. ‘Remarks’ below). Genitalia (fig.
42): Tergum IX with posterior 4 overhanging seg-
ment X in lateral view, posterior margin bilobed with
two blunt lateral angles separated by a obtuse mesal
notch in dorsal view; sternum IX dorsomesal process
slender horizontal, with pointed apex curved dorsad
in lateral view. Segment X dorsal margin slightly con-
cave in lateral view, distal portion narrow; ellipsoidal
and with apex slightly incised in dorsal view. Preanal
appendages large lobate and extended dorsoposteriad
in lateral view. Inferior appendages thumblike and in-
clined dorsad in lateral view, ventrobasal shelf with
short free apical projection; distal arms ellipsoidal in
ventral view, without sharp mesal tooth, mesal mar-
gin with deep narrow curved basolateral notch.
Distribution. — Malaysia: Sarawak.
Remarks. — This species was originally described as a
subspecies of D. morosa. Comparison of the male type
of D. nervosella with that of D. morosa reveals that each
is a distinct species. The modified spur in D. nervosella
is similar to that of D. morosa, but more robust and ac-
tually a mirror image of the other, having apical points
curved in opposite directions. This species also differs
by having the male forewing with a more contrasting
colour pattern, and inferior appendage having mesal
margin with deep curved basolateral notch in ventral
view. Also, D. nervosella is only known from Sarawak,
and D. morosa is only known from the islands of
Mindanao and Basilan of the Philippines.
Dipseudopsis nieuwenhuisi Ulmer, 1909
(figs. 6, 43)
Dipseudopsis nieuwenhuisi Ulmer, 1909: 128-129, fig.4,
Type d*: Borneo Exp., Dr. Nieuwenhuis, L. Dengay,
1.X.1894 (RMNH). — Fischer 1962: 12; Fischer 1972: 6.
Specimens examined. — MALAYSIA: Sarawak, Mt Dulit,
4000ft, moss forest, at light, Oxford Univ. Exp., B. M.
Hobby & A. W. Morre (BMNH): 1d, 22.IX.1932; 5d, 19-
29.X.1932; 18, 17.1X%.1932.
Male. — Body and wings (fig. 6). Head dorsum
with dark brown triangle along anterior margin ex-
tending to middle of posterior margin, posterior
warts and lateral margins yellowish, frontoclypeus
dark brown, antennae yellowish brown, palps dark
brown. Maxillary palps 4.0 mm, proboscis 2.8 mm,
labial palps 1.6 mm. Pronotum dark brown except
130
posterior lateral corners light brown, propleuron light
brown, mesonotum and tegula dark brown, meta-
thorax light brown. Forewing 19-23 mm, dark
brown, veins dark brown with several translucent
stripes between veins, similar to that D. digitata, hav-
ing three translucent areas, basal stripe in thyridial
cell, round marginal spot at arculus and subapical
translucent stripes distal to cord. Front and middle
legs with coxae dark brown, trochanters to apex yello-
wish brown; hind leg mostly yellowish. Modified
spur (fig. 43) 2X as long as adjacent spur and 2x as
thick, with three apical points, one minute point, and
two large points parallel and curved, one large point
subapical. Genitalia (fig. 43): Tergum IX posterior
margin broadly bilobed, having shallow mesal notch
in dorsal view. Segment X originating from relatively
shallow cavity of tergum IX in lateral view, apex
blunt, ventral margin slightly concave; ellipsoidal and
posterior margin broadly bilobed in dorsal view.
Preanal appendages triangular and rounded with dor-
sal margin horizontal, broadly curved and slightly
emarginate in lateral view, ventroposterior margin
broadly curved and strongly inclined dorsad. Inferior
appendages oblong with ventrobasal shelf in lateral
view; angulate with distal arms thumblike in ventral
view, mesal margin with deep narrow curved basola-
teral notch.
Distribution. — Malaysia: Sarawak.
Remarks. — This is the largest Asian species of
Dipseudopsis known. It appears to be closely related to
D. digitata and D. nervosella, but differs by its larger
size and by characteristics of the male modified spur
and genitalia.
Dipseudopsis notata (Fabricius, 1781)
(fig. 44)
Phryganea notata Fabricius, 1781: 390, Type d*: ‘N.
America’ [mislabelled, cf. ‘Remarks’ below], (BMNH). —
Fischer 1962: 13.
Dipseudopsis horni Ulmer, 1915: 46-47, fig. 13. Type series:
SRI LANKA: 26 , [N. Cen. Prov.: Mahl Iluppallama] Maha
Illupalama, Ceylon, Juli 1912, J. C. F. Fryer leg., im
Museum Cambridge [MCZC]; ferner 9 Exemplare (4, 2),
(Cen. Prov.]: Kandy, resp. [E. Prov: Kantalai?]
Kanthaley, resp. [Cen. Prov.: Matale] Matala, Ceylon,
Dr. W. Horn leg., im Deutsch. Entom. Museum Berlin-
Dahlem’ [ZMUH]. — Weidner (1964: 73): “Ceylon,
Matala, Paratypoid’ of D. horni, (ZMUH). — Fischer 1962:
9; Weidner 1964: 73; Fischer 1972: 4. Syn. n.
Specimens examined. — INDIA: Karnataka: 83, Bhadra-
vati (MCZC). 2d , Shimoga, R. Tunga 1865ft (MCZC). Tamil
Nadu: 3d, Tanjore dist. “Thanjavur, Nedungadu (MCZC).
SRI LANKA: N. Prov.: 26, Marichchukkaddi (BMNH). N.
Cen. Prov.: 64, Moradan Maduwa, Wilpattu Natl. Park,
23mi W of Anuradhapura, 80m (MZLU). 1d, Kontalai, 60m
(MZLU). Anuradhapura Dist: 75 89, Wildlife Soc.
Bungalow, Hunuwilagama, Wilpattu Natl. Park, 200ft (-
USNM). 106 109, Irrigation Bungalow, [Padawiya Tank]
‘Padaviya’, 180ft (USNM). 88 29, Padaviya, 180ft (USNM).
Polonnaruwa Dist: 2¢ 79, Pinburettawa, 13mi S
Mannampitiya (USNM). 1d, Polonnaruwa (SOFM). N. W.
Prov.: 16 22, Marai Villu (BMNH). 3¢, 5mi NNE of
Puttalam, 5m (MZLU). S. Prov.: 16, Hikkaduwa, 11mi NW
of Galle, 10m (MZLU). 2d 29, Badagiriya Tank, 6.5mi N
Hambantota, 75ft (CLJw). 16 19, Hambantota (BMNH).
16, Tangalla (MCZC). 18 19, Weligama (BMNH). W.
Prov.: 1068 119, Colombo (BMNH). 5d, 18mi NE of
Colombo, Dambuwa Estate, 30m (MZLU).
Male. — Head dorsum mostly brown, posterior
margin lighter, frontoclypeus, antennae, and palps
brown. Maxillary palps 2.3 mm; proboscis 0.9 mm,
as long as first 24 segments of maxillary palp.
Pronotum brown, slightly lighter in colour than head
and mesonotum. Mesonotum and tegula brown.
Forewing 12-14 mm, mostly brown, similar to that of
D. robustior, with arc of small translucent spots distad
of cord, hyaline spot at m-cu and big spot at arculus.
Legs with coxae brown and distal segments lighter.
Modified spur (fig. 44) slightly longer than adjacent
spur, bifid with two unequal sinuate points directed
distally, longer point almost % as long as spur.
Genitalia (fig. 44): Tergum IX apical % overhanging
segment X in lateral view, posterior margin with
broad obtuse angle in dorsal view; sternum IX meso-
superior process triangular and extended above basal
% of phallus in lateral view. Segment X dorsal margin
slightly concave with blunt apex in lateral view, oval-
shaped almost % as wide as IX in dorsal view. Preanal
appendages trapezoidal with dorsal margin horizontal
in lateral view, ventral margin inclined dorsad and
posterior margin slightly incised. Inferior appendages
angled dorsad in lateral view, dorsal margin concave
and nearly parallel with ventral margin, middle of
ventral margin with obtuse angle, apex rounded; dis-
tal arms diverging in ventral view, lateral margins
with base and apex rounded, mesal margin with squa-
rish basomesal angle, and short apicomesal point.
Distribution. — India: Karnataka, Tamil Nadu; Sri
Lanka.
Remarks. — This species has male modified spur bi-
fid and somewhat similar to that of D. robustior, but
differs by having spur with the shorter point nearly
straight and directed distad. Until now the identity of
D. notata has been enigmatic, because the locality
stated by Fabricius (1781) as ‘America boreali’ was
very doubtful. McLachlan (1864) placed this species
in Dipseudopsis, and suggested that the original loca-
lity information by Fabricius was in error, since at the
time the genus was known only from South Africa
WEAVER & MALICKY: Dipseudopsis from Asia
and China. Our examination of the male type of D.
notata reveals that it is identical to Ulmer’s (1915)
original description of D. horni, only known from
southern India and Sri Lanka, and we therefore con-
cur with McLachlan that the type of D. notata is mis-
labelled.
Dipseudopsis recta Martynov, 1935
(figs. 15, 45)
Dipseudopsis recta Martynov, 1935: 163-164, fig. 65a-b,
Type, INDIA: ‘d. Sanjai River, Chakradharpur, Chota
Nagpur. 8-10.i1.1918. N. Annandale and F. H. Gravely’
(probably NZSI). — Fischer 1962: 14; Higler 192: 80.
Dipseudopsis bombayana Martynov, 1935: 164-165, figs.
GGa-b, 67, Type, INDIA: ‘d. Gokak, Belgaum district,
Bombay Presidency (at light). June 1928 B. Prashad and
H. S. Rao.” [Type (NZSI) in poor condition; not able to
borrow for examination, pers. comm. Dr. K. Rai]. —
Fischer 1962: 5; Higler 1992: 80. Syn. n.
Specimens examined. — INDIA: Andhra Pradesh: 14,
Lamnasinghi Ghats, 75km SW Vishak, 850m, 10.V11.1982,
D. R. Davis (USNM). Bihar: 1d 52, Namkum Ranchi,
675m, 8.XI.61, Ross & Cavagnaro (CASC). Karnataka: 12d
19, Haveri, 2.XII.1961, F. Schmid (Cncı). 16, Jannapura,
22.1.1959, F. Schmid (CNCI). 2d, Kakankote, 10.1.1959, F.
Schmid (cncı). 4d 89, Yellapur, 30.1.1959, F. Schmid
(ENCI). 28, Shimoga, R. Tunga, 1865ft, III-12.VII, P.S.
Nathan (MCZC). Madhya Pradesh: Satanwara, 376 179,
28.XI.1961; 18 39, 29.XI.1961, F. Schmid (CNCI). 166,
Phalghat, 6.11.1959, F. Schmid (CNCI). Maharashtra: 3d
19, (Bombay) Sykes, 5.1.1959, F. Schmid (cncı). Tamil
Nadu: 36, Varaiyattu Tittu, 21.X11.1958, F. Schmid
(ENCI). 26, Maraiyur, 19.XII.1958, F. Schmid (encı). 16,
Sathuparai, 1.X11.1958, F. Schmid (CNCI). NEPAL: 16,
Chitwan Dist., Sauraha S.I. Camp, 31.X.1985, W. N.
Mathis (USNM).
Male. — Head brown, occipital margin light brown,
frontoclypeus light brown except dorsal margin with
dark band, antennae with dorsal portions brown, and
ventral areas light brown, palps light brown, maxil-
lary palps 3.7 mm, proboscis 1.5 mm, almost as long
as first three segments of maxillary palp. Pronotum
brown, hind margin light brown, lateral sides light
brown; mesonotum brown, with tegula and forew-
ings light brown. Forewing 10-12 mm long, faded ar-
ea distal to cord, small hyaline spots at m-cu and ar-
culus. Legs light brown. Modified spur (fig. 45) about
1.5x long as adjacent spur, apical # with crown of
three minute points, having two larger exterior points
spiraling around minute middle point, middle point
slightly curved, directed distad, inconspicuous and
more lightly sclerotized than other points. Genitalia
(fig. 45): IX tergum with apical % overhanging X in
lateral view, posterior margin slightly incised in dorsal
view; IX sternum mesosuperior process with blunt
dorsal extension and slender posterior lobe overhan-
ging basal 4 of phallus in lateral view. Segment X
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
slightly longer than preanal appendage, with lateral
margins parallel and apex deeply bilobed in dorsal
view. Preanal appendages dorsal margin straight in
lateral view, ventral margin curved dorsad, posterior
margin truncate, apicodorsal angle square, ventropo-
sterior angle obtuse. Inferior appendages similar as in
D. triclavata, basal % twice as thick as apical portion in
lateral view, with dorsal margin almost straight, ven-
tral margin having distal % incised, distal % fingerlike
but slightly acuminate; basal shelves separated by V-
shaped notch in ventral view, posterior arms with
blunt apicomesal point, lateral margin straight with
curved apical and basal angles.
Female. — Tergite of abdominal segment VIII (fig.
15) with ventral submarginal recessed groove.
Genitalia (fig. 15).
Distribution. — India: Andhra Pradesh, Bihar,
Karnataka, Madhya Pradesh, Maharashtra, Tamil
Nadu; Nepal.
Remarks. — D. recta is a sibling species to D. tri-
clavata, but differs by having male modified spur with
smaller apical points. The specific differences shown
in the original species descriptions of D. recta and D.
bombayana can be observed by rotating the modified
spur. Hence, D. bombayana is considered to be a jun-
ior synonym of D. recta.
Dipseudopsis robustior robustior Ulmer, 1929 stat. n.
(fig. 46)
Dipseudopsis robustior Ulmer, 1929: 185-186, fig. 22, Type
series: MALAYSIA: West Malaysia: “Material in Coll.
McLachlan: 14, Perak (etwas verletzt).’ Cotype d *: ab-
domen missing, Perak (BMNH). — Fischer 1962: 14.
Dipseudopsis akhila Schmid & Denning, 1979: 243-244, fig.
2, Holotype d*: THAILAND: Chiang Mai Prov., Mae
Ping, Chiang Mai, 1000ft, 10.X1.1964, Peters (CLDD);
Paratypes*, Tha Phra, Khon Kaen. Syn. n.
Dipseudopsis thailandica Schmid & Denning, 1979: 246, fig.
4, Holotype d*: Tha Phra, Khon Kaen, 24.XII.1968,
Knapp; Paratypes*, Nong Pora Kit bamroog, Roi-et, and
Kalasin. Syn. n.
Dipseudopsis junki Marlier, 1979: 1-10, figs. 1-7, Type se-
ries: 25 72, 4 larvae, 1 pupa, Bung Borapet, lake in cen-
tral Thailand, 5.1X.1971, Dr. W. Junk. Syn. n.
Specimens examined. — BURMA: 1d, Rangoon, MCL. coll
(BMNH). CAMBODIA: 66 42, Phnom Penh (USNM, ZMHB).
MALAYSIA: West Malaysia: Kedah: 1 4, Kuala Ketil (BMNH).
43, Bumbong Lima, light trap (USNM). THAILAND: 26 19,
Bangkok (USNM, MCZC). Chiang Mai: 256 162 (USNM);
13 12 (mczc);5d 19, Chiang Mai, Ban-tin-doi, 310m (-
BPBM). 4d, Klong Rang Sit, at light (BMNH, NHMW). 16,
Payao (USNM). 2d, Bung Borapet (USNM). 4d, det. Banks
(1931b) as D. stellata, ‘Penin. Siam’, Patalung, 1.V.1924, I.
H. N. Evans (MCZC, BMNH). 16, Patalung, Paknam
132
Lampam (BMNH). 8¢, Tha Phra, Khon Kaen (CLDD). 76,
Chiang Mai, River Ping (CLHM, ZMUC). 106, Chiang Mai
Zoo, light trap (CLHM). 278, Tap-Tan (CLHM). 1d,
Chantaburi, Khao Soi Dao, 400m (uopy). 16,
Kanchanaburi Dist., Sai Yok Natl. Park, 400m (CLHM). 1d,
Nam Nao Natl. Park (CLHM). 36 , Puk Hieo, Chayapoom (-
CLHM). 5d, Ayutthaya (CLHM). 24 , Bangsaen (CLHM). 1d,
Tham Than Lod Natl. Park (CLHM). 16, Phuket, Kao
Prataew Natl. Park (CLHM). VIETNAM: 1d, Long Xuyen (-
MNHN). 24, Hue (MNHN).
Male. — Head mostly brown, posterior warts light
brown, frontoclypeus with ventral % light brown and
dorsal part brown, antennae mostly yellowish brown
but dorsal surface dark brown, palps yellowish brown,
maxillary palps 2.1 mm, proboscis 1.1 mm, as long as
first three segments of maxillary palp. Pronotum and
tegula light brown, mesonotum dark brown.
Forewing 11-12 mm, similar as in D. collaris, with 5-
7 small translucent spots in circular pattern around
discal cell, having most spots distal to cord. Legs with
coxae dark brown, distal segments yellowish brown.
Modified spur (fig. 46) bifid, with two apical points,
shorter point stout and angular, longer point slender,
semicircular, and distant from other point. Genitalia
(fig. 46): Tergum IX with short apical lobe extending
over X in lateral view, posterior margin triangular in
dorsal view; sternum IX mesosuperior process with
short rounded dorsal lobe and long slender fingerlike
posterior lobe in lateral view. Segment X dorsum in-
clined steeply ventrad toward blunt acuminate apex
in lateral view, apex directed posteriad, having middle
of ventral margin curved ventrad, and apicoventral
margin slanted dorsad, ellipsoidal in dorsal view.
Preanal appendages trapezoidal in lateral view, dorsal
margin straight, ventral margin slanted dorsoposteri-
ad, apicodorsal angle obtuse, apicoventral apex
curved and extended more posteriad than apicodorsal
angle. Inferior appendages with dorsal and ventral
margins parallel and apical % bent dorsad in lateral
view, basal % with thin basal shelf, and apical % trape-
zoidal with apex slighted rounded, distal arms almost
fingerlike in ventral view, lateral margins with apical
and basal ends curved, basoventral shelf short, and
mesal margin with squarish basal angle.
Distribution. — Burma; Cambodia; Malaysia: West
Malaysia; Thailand; Vietnam.
Remarks. — D. robustior is the most common spe-
cies of Dipseudopsis in Thailand, and is somewhat dif-
ficult to distinguish from D. collarisand D. infuscata.
Possibly these species are merely variants of a widely
distributed species.
WEAVER & MALICKY: Dipseudopsis from Asia
a: Big.
Figures 46-50. — Dipseudopsis, A-G, male genitalia: A, lateral; B, preanal appendages, segments IX and X, dorsal; C, inferior ap-
pendages, ventral; D, phallus, lateral; E, phallus, ventral; F, segments IX, X, dorsal; G, ventral. H, see specific descriptions below.
I-J, male modified apical spurs of hind tibia, ventral. Fig. 46, D. robustior Ulmer. Fig. 47, D. robustor andamanensis, n. ssp. 47 I,
modified spur, isolated spur depicted at left. Fig. 48, D. schmidi, n. sp., 48H. left inferior appendage, ventral, variation with ap-
icomesal tooth. Fig. 49, D. spectabilis Banks. Fig. 50, D. stabatensis Malicky & Weaver; 50H, segment IX, dorsal.
133
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Dipseudopsis robustior andamanensis ssp. n.
(fig. 47)
Dipseudopsis infuscata. — Malicky 1984: 214, 216 [misdet.].
Type data. — Holotype d : INDIA: South Andaman Island:
Mongelutonge, 20.XII.1976, Starmühlner (CLHM). —
Paratypes: 18 59, same data (CLHM); 28 159, Bimblton,
22.XI1.1976, Starmühlner (CLHM).
Male. — Head, body, wings and genitalia similar to
D. robustior. Modified spur (fig. 47) with two apical
subequal in length and size.
Remarks. — This subspecies of D. robustior has
unique, pincher-like modified spur (fig. 47) with two
subequal points, but in D. robustior (fig. 46) the api-
cal points of the modified spur are unequal, having a
long slender curved point and a short broad point. It
differs from D. infuscata (fig. 41) which has the mod-
ified spur with a long straight point directed distad
and a shorter curved point. In D. collaris the points
are shorter, only % as long as the spur. This subspecies
represents the only member of the genus known from
the Andaman Islands. All previous records of D. in-
fuscata from South Andaman Island (Malicky 1984)
are now recognized as this subspecies.
Etymology. — This subspecies is named after the
Andaman Islands.
Distribution. — India: South Andaman Island.
Dipseudopsis schmidi sp. n.
(fig. 48)
Type material. — Holotype d: INDIA: Manipur:
Kaiphundai, 20.V.1960, F. Schmid (CNCI). Paratypes: 9d,
same data, F. Schmid (CNCI). 16 , Kambiron, 24.V.1960, F.
Schmid (CNCI). Assam: 6d 29, Garampani Res., 10mi S
Golaghat, 110m, 9.X.61, E. S. Ross & D. Cavagnaro (-
CASC). 1d, Kaziranga, 75m, 26.36N 93.28E, 7-9.V.1976,
Wittmer (CLHM). ‘U.D.M.N.C.H’ [United District of
Mikir and North Cachae Hills]: 1d, Langtrang,
30.IV.1960, F. Schmid (CNCI). BANGLADESH: 1d 29, Lawa
Chera For., Srimangal, 110m, 27.IX.61, E. S. Ross & D.
Cavagnaro (CASC).
Male. — Head dorsum, pronotum and cervical
sclerites light orangish brown, but head slightly dark-
er. Frontoclypeus dark brown except for small oran-
gish brown triangular area between bases of antennae;
genae yellowish brown except ventral sides dark
brown. Scape mostly orangish brown, but apical por-
tion and remainder of antenna dark brown.
Mouthparts, with palps dark brown, maxillary palps
2.5 mm, proboscis 1.5 mm, as long as first four seg-
ments of maxillary palp. Mesonotum and tegula dark
brown. Forewing 11-13 mm, dark brown with indis-
134
tinct translucent spots, large basal triangular translu-
cent spot bordered by Sc, Cu, and first branches of Rs
and M, another translucent spot distal to cord.
Hindwing lighter brown than forewing, with translu-
cent basal triangular area between Sc and Cu,. Legs
with colouration similar as in D. nebulosa, mostly
dark brown, but apical % of front and middle femora
lighter brown, as well as tibiae and tarsi. Hind leg
mostly dark brown, tarsi slightly lighter. Modified
spur (fig. 48) with basal 4 unmodified, distal 4 with
setose pocket at base of straight acuminate apex
pointed distad. Genitalia (fig. 48): Tergum IX broad
and triangular in dorsal view, long with apical 4
overhanging segment X in lateral view. Segment X
broad, extending distad slightly further than inferior
appendages in lateral view, almost twice as long as
broad and somewhat ellipsoidal, with dorsal and ven-
tral margins inclined ventrad but extended posteriad
at apex, apex broadly curved; elongate and cordate in
dorsal view, with lateral margins tapered distad and
apex slightly incised. Preanal appendages broad and
rhomboid in lateral view. Inferior appendages broad
and somewhat rectangular with curved ventral mar-
gin in lateral view, dorsal margin with short broad ba-
sal hump, maximum height about % total length.
Etymology. — Named after its collector Fernand
Schmid.
Distribution. — Bangladesh; India: Assam, Mani-
pur.
Remarks. — This species is easily recognized by hav-
ing the male modified spur acuminate and nearly
straight. The male is somewhat similar to D. nebulo-
sa, but differs by having light areas of forewing with
faded borders and less distinct, and sternum IX with
mesosuperior process not short and globular, but long
and slender, extending above basal % of phallus.
Dipseudopsis spectabilis Banks, 1931
(figs. 9, 49)
Dipseudopsis spectabilis Banks, 1931a: 69, pl. 5: fig. 1, Type
G *: MALAYSIA: Sabah: Sandakan, Baker (MCZC 16421). —
Fischer 1962: 15.
Male. — Head dorsum with anterior portion dark
brown, posterior warts and lateral margins yellowish
brown, frontoclypeus dark brown, genae and anten-
nal scape yellowish brown, but flagellum dark brown,
palps dark brown; maxillary palps 1.8 mm, labial
palps 0.6 mm, proboscis 1.4 mm, longer than first
four segments of maxillary palp. Pronotum, mesono-
tum and tegula dark brown. Wings (fig. 9): Forewing
11 mm, fork I absent, conspicuous pattern of lighter
bands on dark brown background, similar as in D.
flavisignata, with five translucent spots: 1) long slen-
der stripe along anterior margin, # as long as Sc; 2)
small oval-shaped spot at base of Rs; 3) cordate spot
in discal and medial cells; 4) truncate V-shaped band
running basad from subapex to crossvein r-m, then
distal to cord to base of M,, and to submargin
through fork IV including M, 5) long irregular semi-
circular band running from base of anal margin, ta-
pering at m-cu, and widening toward margin at arcu-
lus. Modified spur (fig. 49): trifid, with three apical
points, first point longest, slightly curved at base and
% as long as spur, second point % as long as spur, and
third point minute and adjacent to second point and
located in curved notch between first and second
points. Genitalia (fig. 49): Tergum IX with posterior
margin slightly extended over segment X in lateral
view, posterior margin broadly rounded in dorsal
view; sternum IX mesosuperior process acuminate,
dorsal margin irregular, ventral margin nearly
straight, extending above basal % of phallus. Segment
X dorsal and ventral margins nearly parallel in lateral
view; thumblike with lateral margins parallel in dorsal
view. Preanal appendages trapezoidal in lateral view,
posterior margin slightly incised and lateral side with
ridge parallel to anterior margin. Inferior appendages
heavily sclerotized, fingerlike with dorsal and ventral
margins tapering slightly distad in lateral view, having
short ventrobasal shelf, and dorsal margin with some
minute bumps; squarish in ventral view, with lateral
arms fingerlike with lateral margins converging
slightly distad and mesal margin finely serrate.
Distribution. — Malaysia: Sabah.
Remarks. — This species is only known from the
unique type; it is a sibling species to D. flavisignata
and differs from it in having the male forewing with a
translucent subapical truncated V-shaped band. It al-
so differs by characteristics of the male modified spur
and genitalia.
Dipseudopsis stabatensis Malicky & Weaver, 1988
(fig. 50)
Dipseudopsis stabatensis Malicky & Weaver, 1988: 4-5, fig.
Sa-e, Holotype d*: INDONESIA: Sumatra: Dolok
Merangir, 2-18.V.1971, Diehl (CLHM). Paratype d*:
Stabat 23.V1.1974 (CLJW).
Dipseudopsis morosa. — Banks 1924: 450 (pro parte) [misdet.
syntype 1 4 *: INDONESIA, Kalimantan, ‘Borneo’, Telang,
X.1881 (MCZC)].
Male. — Head dorsum with dark brown median
stripe tapering posteriad, lateral sides and posterior
warts yellowish brown, frontoclypeus dark brown, ge-
nae light brown, antennae light brown, palps light
WEAVER & MALICKY: Dipseudopsis from Asia
brown, maxillary palps 2.4 mm, proboscis 1.2 mm, as
long as first three segments of maxillary palp. Prono-
tum with raised areas yellowish brown, but deep me-
dian furrow and lateral sides dark brown. Mesono-
tum and tegula dark brown. Metanotum light brown.
Forewing 13 mm, brown with translucent apical and
anal spots, hyaline spots at m-cu and arculus. Fore
and mid legs with coxae dark brown and distal seg-
ments yellowish brown, hind leg yellowish brown.
Modified spur (fig. 50) 14x longer than adjacent
spur, apex bifid with two minute curved apical
points, each about % length of spur. Genitalia (fig.
50): Tergum IX posterior margin bilobed, with lobes
separated by squarish mesal notch in dorsal view,
minute portion of posterior margin overhanging seg-
ment X in lateral view; sternum IX mesosuperior pro-
cess similar to that of D. flinti, but slightly slenderer
in lateral view. Segment X thumblike with lateral
sides parallel and sinuate in dorsal view. Preanal ap-
pendages ellipsoidal and extended dorsoposteriad in
lateral view. Inferior appendages capitate with trape-
zoid apex extended dorsad in lateral view; lateral arms
fingerlike with lateral margins straight and parallel in
ventral view, and mesal margins with broad curved
basolateral incisions.
Distribution. — Indonesia: Kalimantan, Sumatra.
Remarks. — This species is similar to D. flinti, but
differs by having inferior appendage slenderer in late-
ral view, and the modified spur with apical points
shorter.
Dipseudopsis tonkinensis Navas, 1921
(fig. 51)
Dipseudopsis tonkinensis Navás, 1921: 79-80, Holotype d *:
VIETNAM: “Tonkin, Chogahn, 1919, P. de Joannis’ (-
MZBS). — Fischer 1962: 16; Fischer 1972: 8.
Specimens examined. — CHINA: Hainan: 1d, Ta Hau,
4.V11.35, J. L. Gressitt (MCZC).
Male. — Head dorsum brown, posterior margin
lighter brown, frontoclypeus brown, antennae brown
but ventral side lighter brown, palps brown, maxillary
palps 2.2 mm, proboscis 1.0 mm, as long as first three
segments of maxillary palp. Pronotum brown, but
posterior margin and lateral sides yellowish brown.
Mesonotum and tegula brown. Forewing 12.5 mm,
faded brown with small hyaline spots at m-cu and ar-
culus. Legs with coxae brown and distal segments yel-
lowish. Modified spur (fig. 51) bifid, with two robust
acuminate points, first point directed obliquely distad
and second point curved mesad. Genitalia (fig. 51):
Tergum IX posterior margin only slightly overhan-
ging segment X in lateral view, posterior margin
135
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
54C
Figures 51-54. — Dipseudopsis, A-G, male genitalia: A, lateral; B, preanal appendages, segments IX and X, dorsal; C, inferior
appendages, ventral; D, phallus, lateral; E, phallus, ventral; F, segments IX, X, dorsal; G, ventral. I-J, male modified spurs ap-
ical spurs of hind tibia, ventral. Fig. 51, D. tonkinensis Navás. Fig. 52, D. triclavata Martynov. Fig. 53, D. varians Ulmer. Fig.
54, D. voluta Ulmer.
136
broad triangular in dorsal view; sternum IX mesos-
uperior process in lateral view with blunt dorsal pro-
jection and acuminate posterior process extended
above basal % of phallus. Segment X dorsal margin
slightly concave and slanted ventrad having blunt
apex in lateral view, ellipsoidal with apex slightly in-
cised in dorsal view. Preanal appendages dorsal mar-
gin straight and ventral margin inclined dorsad distad
with posterior margin broadly incised in lateral view.
Inferior appendages, clavate, being slightly broadened
in lateral view, bent dorsad with oblique angle in
middle, dorsal and ventral margins parallel, apex
truncate; apical arms with apicolateral margins
rounded and slightly broadened in ventral view, api-
comesal angles squarish, mesal margins with square
basolateral notch and square basomesal shelf.
Distribution. — China: Hainan; Vietnam.
Remarks. — This species is similar to D. collaris, but
differs by having male modified spur with apical
points shorter, and by having the male genitalia with
segment X broader in dorsal view.
Dipseudopsis triclavata Martynov, 1935
(fig. 52)
Dipseudopsis triclavata Martynov, 1935: 161-162, figs. 62a-
b, 63a-b, 64. New Lectotype d*: INDIA: Maharashtra:
Khanapur, Belgaum dist., Bombay Presidency, 18-
19.X1.1928, B. P. & H. S. R. (NZsi). Remainder of type
series: 19, same data as lectotype. 1d [doubtful det.],
Meghalaya: The Peak, Shillong, 8,400ft, 25.34N 91.53,
12.X.1914. S. W. Kemp. — Fischer 1962: 16; Higler
1992: 80.
Dipseudopsis morosa. — Schmid 1958: 13, 24, 27, 89, pl. 15:
figs. 12-13 [misdet.].
Specimens examined. — INDIA: Karnataka: 3d,
Uppinangadi (CNCI). 16, Patan (CNCI). 146 49, Shimoga,
R. Tunga, 1865ft (MCZC). 2d 109, Bhadravati (MCZC). 8d
49, Kakankote (CNCI). SRI LANKA: E. Prov.: Amparai Dist.:
1d, Inginiyagala, 250ft (USNM). Cen. Prov.: 1d det. Ulmer
as D. stellata, Peradeniya (BMNH). Kandy Dist.: 118 69,
Hasalaka, Dam Spillway, 300ft (USNM). Matale Dist.: 24
29, Sigiriya, 800ft (USNM). Nuwara Eliya Dist.: 24, Milk
Board Dairy, 4.2mi SW Nuwara Eliya, 6200ft (USNM). N.
Prov.: Vavuniya Dist: 8d 29, Parayanalankulam Irrigation
Canal, 25mi NW of Medawachchiya, 100ft (USNM). N.
Cen. Prov: Anuradhapura Dist: 48 29, Wildlife Soc.
Bungalow, Hunuwilagama, Wilpattu Natl. Park, 200ft (-
USNM). NW. Prov.: Puttulam Dist.: 18, Wilpattu Natl.
Park, 100ft, Tala Wila (USNM). Sabaragamuwa Prov.: 1d,
Niriella (USNM). Ratnapura Dist: 156, Walawe Ganga,
Embilipitiya, 100ft (USNM). S. Prov.: Galle Dist: 1d 19,
Hiniduma (USNM). Hambantota Dist: 1d, Palatupana,
10ft (USNM). Uva Prov: Badulla Dist: 16 19, Dunhinda
Falls, 1300ft (USNM). Monaragala Dist: 9d, Sella
Karagama, Menik Ganga, 150ft (USNM). W. Prov:
Colombo Dist.: 18, Malewana (USNM).
WEAVER & MALICKY: Dipseudopsis from Asia
Male. — Head dorsum brown, except narrow yello-
wish brown posterior margin; frontoclypeus mostly
yellowish brown with brown stripe along dorsal mar-
gin, antennae dorsum brown and ventral sides yello-
wish brown; palps light brown, maxillary palps 2.5
mm, proboscis 1.3 mm, as long as first three segments
of maxillary palp. Pronotum mostly brown, but late-
ral sides yellowish brown, propleuron and cervical
sclerites yellowish brown. Mesonotum and tegula
brown. Forewing 11-14 mm, dark brown with trans-
lucent stigma and hyaline spots at m-cu and arculus.
Fore and mid legs with coxae to mid femora dark
brown and apical % of femora and distal leg segments
light brown; hind leg yellowish brown. Modified spur
(fig. 52) trifid, apex of spur appears truncate, but with
three distinct apical points spiraling in plane perpen-
dicular to longitudinal axis of spur, having one min-
ute point and two longer points, long points each % as
long as spur. Genitalia (fig. 52): Tergum IX with
minute portion of apex overhanging segment X in lat-
eral view, posterior margin truncate with broadly
rounded lateral corners in dorsal view. Sternum IX
mesosuperior process with rounded triangular dorsal
process and smaller posterior lobe extended above ba-
sal # of phallus in lateral view. Segment X dorsal mar-
gin slightly concave and inclined steeply ventrad in
lateral view, apex acuminate with rounded end, ellip-
soidal with apex incised in dorsal view. Preanal ap-
pendages trapezoidal with dorsoposterior apex broad-
ly rounded and dorsal and posterior margins concave
in lateral view. Inferior appendages similar to D. rec-
ta, base 2X as thick as fingerlike apical % in lateral
view, ventral margin of apical portion incised; basal
shelves with broad V-shaped medial notch in ventral
view, apical arms with middle of mesal margin having
oblique incision, lateral apical margin rounded, bas-
olateral margin squarish and apicomesal angle squa-
rish with short blunt tooth.
Distribution. — India: Karnataka, Maharashtra; Sri
Lanka.
Remarks. — This species is closely related to D. rec-
ta, but it differs by having male modified spur with
longer apical points. There has been some confusion
regarding the identity of this species, due to an erro-
neous redescription of ‘D. morosa’ by Schmid
(1958), based on a specimen from Sri Lanka that was
actually D. triclavata. D. morosa is only known from
the southern islands of the Philippines. The male
modified spurs of these species are similar, however
the spur of D. triclavata (fig. 52) differs by being tri-
fid, but that of D. morosa (fig. 39) is bifid. The record
of the male syntype from Shillong, Meghalaya by
Martynov (1935) is doubtful and should be con-
firmed.
197
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Dipseudopsis varians Ulmer, 1929
(fig. 53)
Dipseudopsis varians Ulmer, 1929: 191-192, figs. 28-30,
Type series: MALAYSIA: West Malaysia: Material in Coll.
McLachlan: 36, Perak.’ Type 1 6 *: Perak (BMNH), con-
dition poor, abdomen and hind legs absent, no other type
material available at Mus. (pers. comm. P. C. Barnard). —
Fischer 1962: 16.
Dipseudopsis petersorum Schmid & Denning, 1979: 345, fig.
3a-e, Holotype d, THAILAND: Chiang Mai Prov., east
fork, Mae Ping at junction of small stream, 59 km north
Chiang Mai, 1350ft, at light, 30.XI.1964, W. L. & J. G.
Peters. Syn. n.
Specimens examined. — BURMA: Tenasserim: 164,
Mekane, 90km E Moulmein, 200m (NHRS). MALAYSIA: West
Malaysia: 15 22?, Melaka, det. Ulmer 1905 as Dipseudopsis
sp., de Malacca’ Presqu’ile, 1899, Errington de la Croix et
P. Chapé (MNHN). 1d, Johor: Kota Tinggi (BMNH). 1d,
Negeri Sembilan: Kuala Pilah, at light (BMNH). 36 , Pahang:
Kuala Tahan, at light, (MCZC, BMNH, BPBM). 2d, Kedah:
Bumbong Lima, light trap (USNM). 1d, Selangor: Kuala
Lumpur (BMNH). THAILAND: 1d, Trang (MCZC). 1d 19,
40mi SE Prachuab, 75m (Casc). 1d, det. Denning as D. pe-
tersorum, Krabinburi, Thung Pho, 5.X.1970, Balmer (-
CLDD). 16, Phuket, Tonesai waterfall (SOFM). 16, Khao
Soy Dan (CLHM). 16, Chantaburi, Phliu, 100m (UOP)).
2d, Loei, Phu Rua, 800m (uory). 16, Nakorn Nayok
Prov., Khao Yai Natl. Park, 700m (ZMUC). 236 79,
Chiang Mai Zoo, light trap (CLHM).
Male. — Head, cervical sclerites, and pronotum
orangish brown. Mesonotum, tegula, frontoclypeus,
palps and antennae dark brown, but genae lighter
brown. Maxillary palps 2.4 mm, proboscis 1.3 mm,
as long as first three segments of maxillary palps.
Forewing 12 mm, colouration similar to that of D.
nebulosa but borders of translucent spots more indis-
tinct, dark brown background, large triangular spot
between bases of R, and Cu, and translucent subapi-
cal spot proximal to fork I, similar to bilobed spot of
D. nebulosa but second lobe below R, less distinct.
Legs with coxae dark brown, distal segments light
brown. Modified spur (fig. 53) shorter than adjacent
spur, with modified apical portion % as long as spur,
having spiraling point with tuft of setae in basal pock-
et. Genitalia (fig. 53): Tergum IX with apical % or
more overhanging segment X in lateral view, triangu-
lar with posterior margin rounded in dorsal view;
sternum IX mesosuperior process slender and acumi-
nate in lateral view. Segment X oblong with dorsal
and ventral margins parallel in lateral view, apex
broadly rounded with venter slightly incised; cordate
and broad with lateral sides tapering distad in dorsal
view, apex blunt and slightly incised. Preanal appen-
dages D-shaped with obtuse dorsal angle in lateral
view. Inferior appendages oblong with irregular dor-
sal and ventral margins in lateral view, ventral margin
curved with obtuse basal bump, dorsal margin irregu-
138
lar with conspicuous tooth in middle, distal arms fin-
gerlike in ventral view, with lateral margins diverging
slightly distad and basal shelves broad.
Distribution. — Burma; Malaysia: West Malaysia;
Thailand.
Remarks. — This species is somewhat similar to D.
immaculata, but differs by having male with modified
spur shorter and twisted like a short corkscrew, and
by having the male genitalia with inferior appendages
with a mid dorsal tooth in lateral view and without an
apicomesal tooth in ventral view.
Dipseudopsis voluta Ulmer, 1906
(fig. 54)
Dipseudopsis voluta Ulmer, 1906: 87-88, 110, fig. 89, Type
d *: INDONESIA: Sulawesi, ‘Makassar’, McL. coll, without
abdomen and hind legs (BMNH). — Fischer 1962: 17;
Fischer 1972: 8.
Specimens examined. — INDONESIA: Sulawesi: 1d, with-
out hind legs, ‘Makassar’, McL. coll (BMNH). Tenggara: 16,
Desa Aopa, 50m, 27.X.1989, at light, R. de Jong & J.
Huisman (RMNH). 1d, Moramo, Sg Sena, 50m,
15.X1.1989, at light, R. de Jong & J. Huisman (RMNH).
Male. — Head dorsum mostly dark brown, posteri-
or warts light brown; frontoclypeus brown with dor-
sal portion dark brown, genae light brown, antennal
scape dark dorsad and light brown ventrad. Maxillary
palps 2.3 mm, apical segments 1-3 yellowish brown,
but basal segments 4-5 dark brown; proboscis 1.1
mm, as long as first three segments of maxillary palps,
labial palps 0.5 mm, second and third segments min-
ute. Pronotum mostly brown, posterior margin light
brown with a few darker muscle scars, and propleu-
ron yellowish brown. Mesothorax with notum and te-
gula dark brown, epimeron dark brown, episternum
and sternum yellowish brown. Forewing 14-15 mm,
dark brown with golden setae, and some translucent
spots between R,,, and R,,,, M,,, and M, and hyaline
spots at m-cu and arculus. Front leg with anterior
portion of coxa dark brown, remainder of leg yello-
wish brown, mid and hind legs yellowish brown, but
hind tibia with brown ring at subapical spur.
Modified spur (fig. 54) 2 longer than adjacent spur,
bifid with long subapical point spiraling around long-
er straight point, about % as long as spur. Genitalia
(fig. 54): Tergum IX broad with posterior margin
blunt triangular in dorsal view; sternum IX dorsome-
sal process with blunt posterior lobe in lateral view.
Segment X similar as in D. tonkinensis, but base of
ventral margin nearly straight and apex curved dorsad
in lateral view. Inferior appendages basal % with slen-
der squarish ventral shelf, distal 4 of ventral margin
bent dorsad, dorsal margin slightly concave, apex
broadened slightly and obtuse.
Distribution. — Indonesia: Sulawesi.
Remarks. — This species is easily recognized by hav-
ing the modified spur with a long straight apical point
and another more slender point spiraling completely
around its base. The genitalia of D. voluta are similar
as in D. tonkinensis, but differs by having inferior ap-
pendages with short obtuse subapical mesal tooth in
ventral view, and with apical portion slightly tapered
in lateral view. This species was previously only
known from the male type which lacks the abdomen
and is in rather poor condition. Two male speci-
mens were collected recently, making it possible to
provide a description of the male genitalia for the first
time.
NOMINA DUBIA
Dipseudopsis onychophora Navas, 1935
nomen dubium
Dipseudopsis onychophora Navás, 1935: 104-105, Type d:
INDIA: ‘Lonawla (Bombay), 9-X-1934. H. Benavent, S. J.’
[location of type unknown]. — Fischer 1962: 14; Higler
1992: 80.
Remarks. — The distributional data of other species
of Dipseudopsis in India and the figure provided in the
original description of D. onychophora by Navas
(1935) of the modified spur of the male type, suggests
that this species is probably synonymous with D. #ri-
clavata. However, since we are unable to locate and
examine the type, the status of this species remains
uncertain.
Dipseudopsis orientalis (Navás, 1913)
nomen dubium
Esperona orientalis Navas, 1913: 12-13, fig. 2a-c, Type, sex
unknown: VIETNAM: ‘Hanoi (Tonkin). Un échantillon
envoyé et cédé généreusement pour ma collection par M.
Laboissiere.’ [location of type unknown]. — Fischer 1962:
14.
Remarks. — We were not able to locate and ex-
amine the type of this species, hence its identity re-
mains uncertain. D. orientalis could be synonymous
with one of the three species of Dipseudopsis known
from Vietnam, D. benardi, D. robustior, and D. tonk-
inensis. According to the original species description
(Navas 1913) the forewing of the type of D. orientalis
is 12.4 mm, suggesting that this species is probably
synonymous with either D. robustior or D. tonkine-
nsis.
WEAVER & MALICKY: Dipseudopsis from Asia
POLYCENTROPODIDAE ULMER
Eodipseudopsis Marlier 1959 nomen dubium
Eodipseudopsis Marlier 1959: 117. Type species: Eodipseudo-
psis tomensis Marlier (original designation).
The genus Æodipseudopsis is removed from the
Dipseudopsidae and is provisionally placed in the
Polycentropodidae. Three of its characteristics sup-
port its removal from Dipseudopsidae: 1) Female
genitalia as figured in the original description have
sternum VIII bearing a pair of ventrolateral processes
(Marlier 1957: fig. 10A). 2) The antennae (cf.
Marlier 1959: fig. 48A) are slender with their bases
distant from each other. 3) The long slender lobe of
the sent gland of sternum V (Marlier 1962: fig 10B)
is uncharacteristic of dipseudopsid females examined,
i.e. Dipseudopsis, Hyalopsyche, Phylocentropus.
tree 0
0 Philopotamidae
Hal fF Psychomyiidae
162 Lo Xiphocentronidae
E Ecnomidae
14 = Polycentropodidae
fe CL, Hydropsychidae
17 [TÈ Phylocentropus
List me Hyalopsyche
13= Cai Hyalopsychella
ff m Dipseudopsis
1227 Protodipseudopsis
Lo Limnoecetis
tree 1
m Philopotamidae
pol m; Psychomyiidae
17-=2 Xiphocentronidae
5 Ecnomidae
all, Polycentropodidae
m =: Hydropsychidae
18 =8 Phylocentropus
—16 m’ Hyalopsyche
—14=—=11 Hyalopsychella
C CS Dipseudopsis
ES Protodipseudopsis
9 Limnoecetis
tree 2 ;
m Philopotamidae
F=20 fF Psychomyiidae
ZIE CE; Xiphocentronidae
Li Ecnomidae
ref 4 Polycentropodidae
| L 3 Hydropsychidae
—19 8 Phylocentropus
ul m: Hyalopsychella
L ms Cio Hyalopsyche
El 9 Limnoecetis
nj m Dipseudopsis
12-=7 Protodipseudopsis
Figure 55. Total tree output calculated by Hennig 86 for
Hydropsychoidea, including outgroup and genera of
Dipseudopsidae.
139
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
ES nent
Ecnomidae |
Sen
Fr Hyalopsychella
Mas ee
Limnocoetis
Re
Phylocentropus
Hyalopsyche
Figure 56. Consensus tree of the families of the Hydro-
psychoidea s. str. Weaver, showing phylogenetic relation-
ships of the genera of Dipseudopsidae.
ACKNOWLEDGEMENTS
We are grateful to the following institutions and
curators for transacted loans for material examined:
British Museum of Natural History, London,
Stephen J. Brooks and Peter C. Barnard; Bishop
Museum, Honolulu, Scott E. Miller, Gordon M.
Nishida and Keith Arakaki; California Academy of
Sciences, San Francisco, Wojcieck J. Palawski, Paul
H. Arnaud, Jr., and Vincent F. Lee; Canadian
National Collection, Ottawa, Fernand Schmid; per-
sonal collection, D. G. Denning; Entomological
Laboratory, University of Osaka Prefecture, S.
Moriuti; Instituut voor Taxonomische Zoologie
Amsterdam, L. Botosaneanu; Museum of
Comparative Zoology, Harvard University, David G.
Furth; Muséum National d'Histoire Naturelle, Paris,
Drs. Jean Legrand, and Jeanine Casewitz-Weulersse;
Museu de Zoologia, Barcelona, O. Escola; Museum
for Zoology, University of Lund, L. Cederholm;
Naturhistorisches Museum Wien, A. Kaltenbach and
U. Aspöck; Nationaal Natuurhistorisch Museum,
Leiden, Jan van Tol and Jolanda Huisman;
Naturhistoriska Riksmuseet, Stockholm, G. Hallin;
Zoological Institute of the Russian Academy of
Sciences, St. Petersburg, Vladimir D. Ivanov and
Lidija A. Zhiltsova; Smithsonian Institution, United
States National Museum of Natural History,
Washington, Oliver S. Flint, Jr; Zoologisches
Museum der Humboldt-Universität Berlin, W. Mey;
Zoologisk Museum Kobenhavn, ©. Karsholt;
Zoological Museum Sofia, K. Kumanski and P.
Beron; Zoological Survey of India, Calcutta, K. Rai
and S. K. Ghosh; and Zoologische Staatssammlung
Miinchen, E. Burmeister.
We are also grateful to the following colleagues for
giving valuable material from their personal collec-
tions: M. Allen, P. Chantaramongkol, E. Diehl, J. A.
W. Lucas, R. Miiller, J. Olah, V. Puthz, W. Speidel,
140
F. Starmiihlner, Tian Li-xin, and W. Wittmer. Also,
we thank Arturs Neboiss, Museum of Victoria, for
providing an editorial review of the manuscript and
transferring borrowed material from RMNH, John F.
Burger, University of New Hampshire, for reviewing
the manuscript, and Matthew Kelly for most of the il-
lustrations. The support of the National Science
Foundation (grant BSR8907401) is gratefully ac-
knowledged.
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Tijdschrift voor Entomologie
Volume 137, no. |
Articles
27
57
75
87
95
©
A. J. de Boer
The taxonomy and biogeography of the loriae group of the genus Baeturia Stal,
1866 (Homoptera, Tibicinidae).
J. Patocka
Die Puppen der Spanner Mitteleuropas (Lepidoptera, Geometridae):
Charakteristik und Bestimmungstabelle der Gattungen.
J. T. Polhemus & D. A. Polhemus
Four new genera of Microveliinae (Heteroptera) from New Guinea.
R. Rozkosny & D. Kovac
Adults and larvae of two Ptecticus Loew from Peninsular Malaysia (Diptera,
Stratiomyidae).
J. van Tol
The Odonata of Sulawesi and adjacent islands. Part 3. The genus Macromia
Rambur (Corduliidae).
J. S. Weaver & H. Malicky
The genus Dipseudopsis Walker from Asia (Trichoptera: Dipseudopsidae).
Nederlandse Entomologische Vereniging, Amsterdam
Published 15 July 1994 ISSN 0040-7496
zi
Cai
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" Marburg, Germany &° The Natural History Museum, London, UK
Re VP ROEMENIE S OUITIERNIPAEAEARCTMIC AND
PAEBEOTROBICHETEAERIOPRPERGENUS
HENGCHUNIA VILBASTE (HOMOPTERA,
CIGDERTIDAE)
SMITHSON TA
APR 1 4 1995 \
LIBRARIES
Asche, M. & M. D. Webb, 1994. Review of thé southern Palaearctic and Palaéotropical leaf-
hopper genus Hengchunia Vilbaste (Homoptera, Cicadellidae). — Tijdschrift voor Entomologie
137: 143-154, figs. 1-63 (Issn 0040-7496). Published 15 December 1994.
The leafhopper genus Pseupalus Remane & Asche (Deltocephalinae: Paralimnini) is synonym-
ized with Hengchunia Vilbaste syn. n. Four new species are described: H. helleri sp. n. from
Cameroun and Zaire, H. indica sp. n. from India, H. pakistanica sp. n. from Pakistan and H.
javana sp. n. from Indonesia: Java. Hengchunia now contains nine species which are probably
all associated with the grass genus /mperata. The zoogeographical distribution of the species and
their potential phylogenetic relationships are briefly discussed. Additions and amendments to
Webb & Heller’s (1990) check-list of Afrotropical and Oriental Paralimnini are made and in-
clude Paralimnellus Emeljanov (1972), oldest available name for Bubulcus Dlabola (1961) (jun-
ior homonym of Bubulcus Bonaparte 1854, Aves) and Paralimnellus vittata (Matsumura)
comb. n. from Henschia.
Correspondence: Dr Manfred Asche, Am Schwanhof 2A, D-35037 Marburg, Germany.
Key words. — Homoptera; Auchenorrhyncha; Cicadellidae; Paralimnini; Hengchunia;
Pseupalus; review; new species; South Palaearctic and Palaeotropical Region.
This paper concerns a group of paralimnine leaf-
hoppers (Cicadellidae: Deltocephalinae) apparently
associated with the Palacotropical grass /mperata, es-
pecially the species 1. cylindrica. This association was
reported by Remane & Asche (1980) who erected the
genus Pseupalus for a single paralimnine species, P.
graecanarus from the Canary Islands (type locality)
and Greece. Recently, Pseupalus and related taxa were
reviewed by Webb & Heller (1990) who added three
more species: P. gaiseri from Ivory Coast, P. digitus
from Thailand, and P. murtus from the Philippines:
Luzon. Webb & Heller (l.c.) discussed the validity of
the tribe Paralimnini sensu Emeljanov (1962),
Remane & Asche (1980), Ossiannilsson (1983), and
Emeljanov & Kirilova (1989) and Hamilton’s
‘Deltocephalina’ (Hamilton 1975), and confirmed
the tribal placement of Pseupalus. In addition, Webb
& Heller (l.c.) provided a check-list of Afrotropical
and Oriental paralimnine genera. Since then, the sec-
ond author has examined specimens of a species from
Taiwan, described as Thamnotettix koshunensis
Matsumura, for which Vilbaste (1969) had estab-
lished the genus Hengchunia. A comparison revealed
not only the placement of Hengchunia in the Oriental
Paralimnini, but also its generic synonymy with
Pseupalus. Moreover, four new species of this genus
were discovered in samples from Africa, India, and
Indonesia, described below, together with an uniden-
tifiable female specimen from Australia. The genus
Hengchunia now contains nine species: one in the
Southwestern Palaearctic, two in the Afrotropical
Region, and 6 in the Oriental Region.
The following amendments and additions apply to
the Afrotropical and Oriental paralimnine check-list
(Webb & Heller 1990), noted above: Paralimnellus
Emeljanov (1972) (type species: cingulatus Dlabola)
oldest available name for Bubulcus Dlabola (1961)
(type species: cingulatus Dlabola) (junior homonym
of Bubulcus Bonaparte, 1854, Aves); Paralimnellus
vittata Matsumura comb. n. (Henschia); Pteropyx ac-
utus (Singh) (Jassargus) Emeljanov, 1972: 66;
Pteropyx maculatus Emeljanov, 1972: 67 (Mali);
Khasia prima Rao, 1989: 82 (India).
Depositories
Abbreviations for depositories of material used in
this paper: M. Asche & H. Hoch, Marburg, private
collection (AH); Agricultural University Wageningen,
The Netherlands (auwn); The Natural History
Museum, London, United Kingdom (BMNH); Musée
143
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 1-10. Hengchunia gaiseri (Webb & Heller), paratype 3 from Ivory Coast; Figs. 6, 10: d from Sierra Leone. — 1, male
pygophore, left lateral view; 2, left subgenital plate, dorsal view; 3, same, dorsolateral view; 4, subgenital plates, ventral view;
5, 6, left style, dorsal view; 7, aedeagus and connective, ventrocaudal view; 8, aedeagus, left lateral view; 9, 10, same, left la-
teral view, slightly bent to left side. Scales: 0.1 mm.
144
Royal de l'Afrique Centrale, Tervuren, Belgium
(MRAC); Institut Royal des Sciences Naturelles de
Belgique, Brussels, Belgium (tsNB); Entomological
Institute, Hokkaido University, Sapporo, Japan (EI-
HU); Staatliches Museum fiir Naturkunde Stuttgart,
Germany (smns); University of Agricultural Sciences,
Bangalore, India (UASB).
‘TAXONOMY
Hengchunia V ilbaste
Hengchunia Vilbaste, 1969: 8. — Type species by original de-
signation: Thamnotettix koshunensis Matsumura.
Pseupalus Remane & Asche, 1980: 88, syn. n. - Type species
by original designation: Pseupalus graecanarus Remane &
Asche.
For relationship and diagnosis of Hengchunia see
the corresponding paragraphs for Pseupalus in Webb
& Heller (1990: 2, 3).
Key to the species of Hengchunia (males)
(after Webb & Heller 1990, modified and
supplemented)
Preatrium of aedeagus with a pair of long and
slemdemprocesses((e 2. Mesi 9 nno D
— Preatrium of aedeagus without a pair of processes
(ie FOR ione H. pakistanica sp. n.
2. Aedeagal shaft with four spinose processes apical-
ya (EE) Nocona eben H. koshunensis (Matsumura)
— Aedeagal shaft with two or less spinose processes
apically or subapically (e.g., fig. 7) ................... 3
3. Aedeagal shaft with a single spinose process or lat-
CAM IAN Eesti 4
— Aedeagal shaft with two spinose processes apical-
rollio ne 5
4. Aedeagal shaft with a horn-shaped subapical pro-
cess directed ventrad, dorsal side of shaft at mid-
length with a bifurcate process and shortly basad
of it with a single short process (figs. 32-34) …
Rn, H. indica sp.n.
— Aedeagal shaft with a subapical flange on each
side, devoid of spinose processes (see Webb &
Elle ISSO fe Bree nn...
oe H. graecanarus (Remane & Asche)
5. Subgenital plates with distal margin slightly con-
cave (fig. 14); aedeagal shaft medially on dorsal
side with a group of 3 short spines (fig. 18) … 6
— Subgenital plates with distal margin convex (e.g.,
figs. 2, 22); aedeagal shaft medially on dorsal side
devoid of short spines (e.g., figs. 8, 26, 61) …. 8
7. Aedeagal shaft with apical processes almost di-
rected laterad; mediodorsal processes compara-
tively distant from each other; style almost evenly
tapering to apex without a subapical lobe (see
ASCHE & WEBB: Review of Hengchunia
Webbrögkleller 1990 Oros MENE) Nn
EEN H. digitus Webb & Heller
— Aedeagal shaft with apical processes directed ven-
trobasad; mediodorsal processes near to each oth-
er (fig. 18); style with a subapical lobe (fig. 16) …
N PAR RTS Ee A. murtus Webb & Heller
8. Aedeagal shaft with processes apically, directed
dorsocaudad, not reflected (figs. 60-62) ............
RR dna H. javana sp. n.
— Aedeagal shaft with processes subapically or me-
dially, directed laterobasad (e.g., figs. 7, 25) … 9
9. Aedeagal shaft with processes located subapically
(fel) re eran H. gaiseri Webb & Heller
— Aedeagal shaft with processes located mediolate-
ralu(tiosı 25126) REP RER H. helleri sp. n.
1. Hengchunia koshunensis (Matsumura)
(fig. 60)
Thamnotettix koshunensis Matsumura, 1914: 178. Lectotype
3, Taiwan (Eru), here designated [examined].
Hengchunia koshunensis: Vilbaste 1969: 8, fig. 6: A-J.
Material examined. — Lectotype d, ‘Formosa/Mat-
sumura and ‘5.vii.1906/Koshun’ on reverse; 19, 1 ? (sex
unknown). — Paralectotypes, originally on same mount as
lectotype (re-mounted) (EIHU).
TG
S
[LA
Fig. 11. Hengchunia murtus (Webb & Heller), d from
Sulawesi. — 11, habitus. Scale: 1.0 mm.
145
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 12-1 92 Hengchunia murtus (Webb & Heller), 5 from Sulawesi. — 12, male genitalia, ventral view; 13, same, left lateral
view; 14, left subgenital plate, dorsal view; 15, same, dorsolateral view; 16, left style, dorsal view; 17, aedeagus and connecti-
ve, ventrocaudal view; 18, aedeagus, dorsal view; 19, same, left lateral view. Scales: 0.1 mm.
146
Remarks. — H. koshunensis can be separated from
the other Hengchunia species by the crown-shaped
apex of the aedeagal shaft consisting of two pairs of
spines: a pair of longer spines directed ventrad and a
pair of shorter spines directed dorsad (not vice versa
as noted in Vilbaste 1969: 8).
Distribution. — This species is only known from
Taiwan (Matsumura 1914; Vilbaste 1969).
2. Hengchunia graecanarus (Remane & Asche)
comb. n.
Pseupalus graecanarus Remane & Asche, 1980: 90, fig. 16 a-
I. Holotype 3, Canary Is, (Remane coll.) [examined]. —
Webb & Heller 1990: 5, fig. 13. .
Material examined (additional to the type material of
Remane & Asche 1980). — 24, 59, S Greece,
Peloponnessos, Skala nr Gythion, on /mperata cylindrica,
6.x.1981, M. Asche & H. Hoch. - 9d, 269, NW Greece,
Corfu I., Aghios Stephanos, coastal biotopes with /mperata
cylindrica, 14.x.1981, M. Asche & H. Hoch. - 3d, 19, NW
Greece, 27 km S. Igoumenitsa, S. Kastri, on /mperata cylin-
drica, 10.viii.1983, M. Asche & H. Hoch (An).
Remarks. — Within the genus, A. graecanarus dis-
plays a unique configuration of the aedeagus with
subapical flanges at the shaft instead of spinose pro-
cesses.
Distribution. — This species is known from the
type-locality in the Canary Islands: Gran Canaria,
and from several localities in Greece (Remane &
Asche 1980). According to the distribution of its host
plant, /mperata cylindrica, a wider distribution at least
in the Mediterranean Region is possible.
3. Hengchunia digitus (Webb & Heller) comb. n.
Pseupalus digitus Webb & Heller, 1990: 4, figs. 14-18.
Holotype d, Thailand (BMNH) [examined].
Remarks. — No material additional to the type ma-
terial has been examined.
H. digitus closely resembles H. murtus in the shape
of the subgenital plates (including the toothed area)
and aedeagus but the aedeagal shaft is straighter and
has the three mediodorsal spines less closely adjacent
and the style apex narrower.
Distribution. — This species has been reported on-
ly from Thailand (Webb & Heller 1990).
4. Hengchunia gaiseri (Webb & Heller) comb. n.
(figs. 1-10)
Pseupalus gaiseri Webb & Heller, 1990: 4, figs. 1-12.
Holotype d, Ivory Coast (smns) [examined].
ASCHE & WEBB: Review of Hengchunia
Remarks. — No material additional to the type ma-
terial has been examined.
H. gaiseri is similar (and probably closely related)
to H. helleri (see below). It differs from this species in
the distally more pointed subgenital plates and in the
position of the lateral spinose aedeagal processes
which are located subapically in A. gaiseri and in
midlength of shaft in A. helleri.
Distribution. — Although this species was described
from Ivory Coast it is likely that a similar male spec-
imen from Sierra Leone is conspecific (Webb &
Heller 1990: 4).
5. Hengchunia murtus (Webb & Heller) comb. n.
(figs. 11-19)
Pseupalus murtus Webb & Heller, 1990: 6. Holotype 6,
Philippines (BMNH) [examined].
Material examined (additional to the type material of
Webb & Heller 1990). — Indonesia: Sulawesi: 16, Sulawesi
Utara, Dumoga Bone National Park, rainforest near base-
camp Toraut, 19.x.1985; 1d, ibid., banks of Toraut river;
16, ibid, light trap near basecamp, 7.xii.1985; 1d, 29, ib-
id, trail to Tumpah river, 23.xi.1985; 1, ibid., trail to
Tumpah river uphill to ca. 400 m, 20.x.1985; 19, ibid.
banks of Tumpah river; 16, Molosso I., opposite Lolok
(N.-coast), 11.xi.1985; all: on Jmperata grass, Project
Wallace Expedition, M. Asche & H. Hoch (AH, BMNH).
Remarks. — H. murtus closely resembles H. digitus
in the shape of the male genitalia (see under H. digi-
tus).
Distribution. — This species was originally descri-
bed from Luzon I. in the Philippines (Webb & Heller
1990). Its occurrence in Sulawesi extends its range to
the south.
6. Hengchunia helleri sp. n.
(figs. 20-26)
Type material. — Holotype d: Zaire, Lubumbasha
(= Elisabethville), 30.11.1939, at light, H.-J. Brédo (1s). —
Paratypes: Zaire: 198, 439, 2 ? (sex unknown), same data
as holotype (various dates) (ISNB, MRAC, BMNH); 36, 39,
Libenge, Savane Liki-Bembe, 23-28.ii.1948 (one without
date) (R. Cremer & M. Neuman, (tsNB); 19, Kivu,
Kinchassa, Waelbroeck, 24.ii and 3.iv.1899 (isNB); 16,19,
Maka, Lualaba, 25.1.1939, H.-J. Bredo (tsNB). - Cameroun:
14, Bertoua, 14.ix.1960, F.J. Breteler (AUWN).
Description. — Habitus, bodily proportions and
colouration as in generic description (see Webb &
Heller 1990).
Male genitalia: Subgenital plate (figs. 20-23) distal-
ly broadly rounded; toothed area rather long, extend-
ing from apex to about half of the length of the sub-
genital plate mediobasad, apical tooth and basal teeth
147
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 20-26. Hengchunia helleri sp.n., paratype d from Cameroun. — 20, male genitalia, ventral view; 21, same, left lateral
view; 22, left subgenital plate, dorsal view; 23, same, dorsolateral view; 24, left style, dorsal view; 25, aedeagus and connecti-
ve, ventrocaudal view; 26, aedeagus, left lateral view. Scales: 0.1 mm.
148
ASCHE & Wess: Review of Hengchunia
Figs. 27-35. Hengchunia indica sp.n., holotype & from India. — 27, male pygophore, left lateral view; 28, left subgenital pla-
te, dorsal view; 29, same, dorsolateral view; 30, subgenital plates, ventral view; 31, left style, dorsal view; 32, aedeagus and
connective, ventrocaudal view; 33, aedeagus, left lateral view; 34, same, dorsal view; 35, apex of aedeagus, left dorsocaudal
view. Scales: 0.1 mm.
strongest. Style (fig. 24) with short inner and long
outer apophysis, the latter continuously tapering to
apex, distal margin with about 7 tooth-like projec-
tions. Aedeagus (figs. 25, 26) with shaft short, evenly
curved dorsad, on both sides at midlength a spinose
process which is directed basad; phallotreme apically,
exposed to the ventrocaudal side; processes of preatri-
um long and slender, straight.
Length: d: 3.5-3.6 mm, 2: 3.8 mm.
Remarks. — H. helleri is similar (and probably
closely related) to H. gaiseri; it can be distinguished
mainly by the shape of the subgenital plates which are
apically less pointed than in H. gaïseri and which bear
a more extended toothed area, as well as by the posi-
tion of the aedeagal processes which arise at mid-
length of the shaft (not subapically as in A. gaiseri).
Distribution. — The occurrence of H. helleri in
Zaire and Cameroun may indicate a wider distribu-
tion, at least in Central- and West Africa.
149
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 36-39. Hengchunia pakistanica sp.n., holotype d from NE Pakistan. — 36, head and thorax, dorsal view; 37, head, fron-
tal view; 38, left tegmen; 39, left hind wing. Scales: 0.5 mm.
7. Hengchunia indica sp. n.
(figs. 27-35)
Type material. — Holotype d: India, Bihar, Pusa,
14.v.1931, T. B. Fletcher (BMNH).
Description. — Habitus, bodily proportions, and
colouration as in generic description (see Webb &
Heller 1990).
Male genitalia: Subgenital plate (figs. 27-30) rather
stout with distal margin very little convex, almost
truncate; toothed area with a triangular portion at
margin and an arm-like extension directed mediobas-
ad. Style (fig. 31) comparatively short and stout, de-
void of a mediodistal apophysis, subapically with a
short lobe-like projection. Aedeagus (figs. 32-35)
with shaft evenly curved dorsad; ventrobasad of the
apical phallotreme with a gently curved horn-shaped
process which arises on a broad base; medially on dor-
sal side of shaft a longer, apically bifurcate process,
and basad of it a shorter single process; processes of
150
preatrium slender, slightly curved ventrad.
Length: 3: 3.0 mm.
Remarks. — Within the genus, H. indica is most
similar to H. pakistanica (see below) with which it
shares the short and distally truncate subgenital plates
and a single horn-shaped aedeagal process basad of
the phallotreme on the ventral side. However, it dif-
fers considerably from this species in the shape of the
toothed area of the subgenital plates and in details of
the aedeagus, especially by the presence of paired pro-
cesses of the preatrium which are entirely missing
(probably reduced) in H. pakistanica.
Distribution. — Only known from the Bihar-
Region in North India.
8. Hengchunia pakistanica sp. n.
(figs. 36-50)
Type material. — Holotype d : Pakistan, Gujrat (= Guja-
ASCHE & WEBB: Review of Hengchunia
Figs. 40-50. Hengchunia pakistanica sp.n., holotype d from NE Pakistan; Figs. 45, 48: paratype d from SE Pakistan. — 40,
male genitalia, ventrocaudal view; 41, same, left lateral view; 42, left subgenital plate, dorsal view; 43, same, dorsolateral view;
44, 45, left style, dorsal view; 46, aedeagus and connective, ventrocaudal view; 47, 48, aedeagus, left lateral view; 49, same,
dorsal view; 50, apex of aedeagus, left dorsocaudal view. Scales: 0.1 mm.
1541
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 51-54. Hengchunia javana sp.n., holotype d from Java. — 51, head and thorax, dorsal view; 52, head, frontal view; 53,
left tegmen; 54, left hind wing. Scales: 0.5 mm.
rat), Nausari, 16.1.1981, C.A. Viraktamath (BMNH). -
Paratypes: 42, same data as holotype (UASB, BMNH); 1d,
12, Pakistan, Hyderabad, Patanduru, vii-ix.1980, Bernays
& Woodhead (BMNH).
Description. — Habitus, bodily proportions, and
colouration as in generic description (see Webb &
Heller 1990).
Male genitalia: Subgenital plate (figs. 40- 43)
short, distally truncate, toothed area separated in an
apical and a median field of teeth which are connect-
ed by a ridge. Style (figs. 44, 45) distally with a short
median edge and a long outer apophysis with a short
subapical lobe. Aedeagus (figs. 46-50) with shaft
short, slightly curved dorsad; a single long, horn-
shaped, ventrobasad directed process arising basad of
phallotreme on the ventral side; shaft submedially
with a pair of lateral processes; preatrium devoid of el-
ongate paired processes.
Length: d : 3.0 mm, 9: 3.1-3.3 mm.
152
Remarks. — H. pakistanica can readily be distin-
guished from all other congeners by the lack of pro-
cesses of the aedeagal preatrium. The aedeagal shaft
with its single horn-shaped process on the ventral side
is similar to that of H. indica but it differs in other
genital characters, as noted above. Slight differences
could by found in the genital structures (especially ae-
deagal spines and styles) between the males from
North and South Pakistan but they seem too small to
warrant species or even subspecific status.
Distribution. — Only known from the Gujrat-
Region in the North-East and from the Sind-Region
in the South-East of Pakistan.
9. Hengchunia javana sp. n.
(figs. 51-62)
Type material. — Holotype d : Indonesia: Java, Bogor, on
Imperata, 24.xi.1989, M.R. Wilson (BMNH). - Paratype:
19, same data as holotype (BMNH).
ASCHE & WEBB: Review of Hengchunia
N
Figs. 55-63. Hengchunia species. — 55-62. Hengchunia javana sp.n., holotype d from Java; 55, male genitalia, ventrocaudal
view; 56, same, left lateral view; 57, left subgenital plate, dorsal view; 58, same, dorsolateral view; 59, left style, dorsal view;
60, aedeagus and connective, ventrocaudal view; 61, aedeagus, left lateral view; 62, apex of aedeagus, ventrocaudal view.
Scales: 0.1 mm. 63; H. koshunensis (Matsumura), aedeagus, redrawn and modified from Vilbaste, 1969, Fig. 6 E.
153)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Description. — Habitus, bodily proportions, and
colouration as in generic description (see Webb &
Heller 1990).
Male genitalia: Subgenital plate (figs. 55-58) distal-
ly convex, toothed area devided in an apical and a me-
dian portion, both portions with prominent teeth.
Style (fig. 59) with a short inner edge and an elongate
outer apophysis with undulate margin. Aedeagus
(figs. 60-62) with shaft short, in middle almost rec-
tangularly bent dorsad; phallotreme apically, dorsally
on each side flanked by a pair of short, fork-like spi-
nose processes directed dorsocaudad; paired processes
of the preatrium conspicuously surpassing the aedea-
gal shaft, almost straight.
Length: d: 3.4 mm; 2: 3.6 mm.
Remarks. — Within the genus, H. javana is rather
isolated. It can readily be separated from the other
congeners by the configuration of the aedeagal shaft
with its fork-like, dorsocaudad directed (not reflect-
ed) spinose processes at apex.
Distribution. — Only known from Bogor in West
Java.
10. Hengchunia sp.
We have examined one female from Australia,
Queensland, Cairns (ridge at Edge Hill, rainforest),
collected on /mperata grass, on 10.vi.1987 by M.
Asche & H. Hoch (an). Externally, this specimen re-
sembles H. murtus, but its specific identity can only
be clarified after a corresponding male becomes avail-
able; however, this finding extends the geographic
range of the genus Hengchunia to NE Australia.
ACKNOWLEDGEMENTS
For the loan of material we would like to thank the
following: Dr. C. Viraktamath (vas), Dr. P.W.F. de
Vrijer (AUWN), Dr. P. Grootaert (1snB), Dr. H. André
(MRAC) and Dr. S. Takagi (Eru). We also wish to
thank Dr. H. Hoch, Museum fiir Naturkunde,
Berlin, Germany, and Dr. M. R. Wilson, National
Museum of Wales, Cardiff, UK, for helpful com-
ments on the manuscript.
154
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dinea: lassidae). — Insecta Matsumurana, Supplement 6.
12pp.
Received: 15 February 1994
Accepted: 28 April 1994
D. B. BAKER
Hope Entomological Collections, University Museum, Oxford, United Kingdom.
A NEW GENUS OF NOMADINE BEES FROM
NORTH AFRICA (HYMENOPTERA: APOIDEA,
ANTHORHORIDAE)
Baker, D. B., 1994. A new genus of nomadine bees from North Africa (Hymenoptera:
Apoidea, Anthoporidae). — Tijdschrift voor Entomologie 137: 155-159, figs. 1-4. [rss 0040-
7496]. Published 15 December 1994.
Aethammobates prionogaster gen. et sp. n. Anthophoridae: Nomadinae) is described from a
unique male collected in Egypt. The new genus is referred to the Holcopasitini and its relation-
ships with other holcopasitine bees are discussed.
D.B. Baker, Hope Entomological Collections, University Museum, Oxford ox] 3pw, U.K.
Keywords. — Hymenoptera; Anthophoridae: Holcopasitini; Aethammobates prionogaster, new
genus; new species; North Africa.
Among a number of epeoline bees received many
years ago from the late Prof. Dr. H. Priesner was a
single male of a strange nomadine bee that could not
be referred to any described genus. A description and
figures were prepared, and the specimen was then set
aside in the expectation that further specimens, in-
cluding possibly the female, would sooner or later
come to light. After more than 40 years, this expecta-
tion has not been fulfilled, although the locality
where the bee was obtained (Gebel el Asfar, near
Cairo) was one much visited by both pre-War and
more recent local collectors. Since the locality is now
being devastated by construction works (C.G. Roche,
in litt., 6 September, 1992), making the recovery of
further material less likely, and since also the system-
atics of the Nomadinae have recently come under re-
view by several authors, notably Alexander (1990)
and Roig-Alsina (1991), it seems desirable to place
the new genus and species on record.
Aethammobates gen. nov.
Description. — Head in frontal aspect transverse,
the eyes rather small, separated by 1.2 times their
length, their inner margins convex, subparallel; clype-
us short, carinate laterally along epistomal suture,
paraocular area adjacent to carina slightly concave
and narrowly impunctate; labrum elongate (length to
breadth ration 1.44 : 1), apically entire, lacking discal
tubercles; antennal sockets at mid-level of eyes; inter-
antennal carina divided, forming a V-shaped protub-
erance, the open end of the V upward; facial foveae
absent; vertex longer than inter-ocellar distance; pre-
occipital ridge carinate; malar area rudimentary.
Antennae 13-segmented, scape long (length to
breadth ratio 3 : 1); pedicel free; first flagellar segment
long (length to breadth ratio 3 : 1), equal to segments
2+3; segments 2-10 passing from transverse to quad-
rate. Mandibles simple, with weak basal protuberance
opposed to lateral termination of clypeal carina; only
posterior articulation in contact with eye. Stipital
comb absent; maxillary palpi 5-segmented; blade of
galea moderately, uniformly sclerotized, acuminate,
extreme apex very narrowly rounded.
Mesosoma broad, dorso-ventrally compressed.
Pronotum with well developed dorsal surface limited
anteriorly by an uninterrupted carina; anterior surface
broad, flat, rounded abruptly into collum.
Mesoscutum little convex; parapsidal lines well
marked, about as long as tegula; axillae not protu-
berant; scutellum extending laterad slightly beyond
axillae, with distinct dorsal and posterior vertical sur-
faces separated by a weak, irregular (interrupted by
punctation) carina, slightly overhanging subvertical,
coplanar surfaces of metanotum and propodeum;
omaulus continuous across mesepisternum, forming
prominent angles on either side of a median, semicir-
cular inflexion; mesepisternum anterior to intermed-
iate coxae strongly transverse, transversely concave.
Metanotum with prominent lateral lobes. Wings (fig.
4) short, forewing about 0.6 body length. Forewing
bare basally, becoming pubescent apically, especially
beyond cells; marginal cell broad, rather broadly
rounded at apex, apex separated from wing-tip by less
than cell length; distal abscissa of Rs absent; two sub-
marginal cells, the second much shorter than the first,
the first receiving 1 m-cu near its apex, the second 2
m-cu beyond the middle. Hindwing with jugal lobe
155
TiyDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
very short; second abscissa of M + Cu about 1,5 times
longer than cu-v, not quite half as long as M. Legs of
normal proportions. Anterior coxae proximate, trans-
verse, the trochanters widely separated; tarsus sub-bi-
pectinate, basitarsus and segments two to four apical-
ly each with a few strong setae on either side;
basitarsus flattened and slightly expanded apicad.
Intermediate coxae widely separated, short, much
shorter than distance from their summits to posterior
wing bases; femur broad; tibia without anterior setose
area, coarsely spiculate. Posterior coxae large, broad,
inwardly weakly, externally strongly carinate; tibiae
coarsely spiculate. Arolia present, normally devel-
oped.
Metasoma broad, depressed, greatest width at apex
of segment 2. Tergum 1 with vertical anterior and
horizontal dorsal surfaces separated by a deeply arcu-
ate carina, in dorsal aspect with prominent lateral an-
gles; marginal areas of terga abruptly and progressiv-
ely more strongly depressed caudad, the margins of
the discal areas passing from subserrate on tergum 1
to strongly denticulate on terga 4-6; pygidial plate
strong, rostriform, rounded apically, not constricted
basally, considerably exceeding apical margin of seg-
ment; sternum 6 laterally dentate.
Vestiture generally squamiform, on the terga form-
ing basal and marginal fasciae, not maculae; simple,
erect hairs largely confined to mandibles and disc of
labrum; anterior angles of malar areas not penicillate;
eyes bare; sterna without subapical fimbriae of modi-
fied hairs, but sterna 5 and 6 mesially with fine,
dense, erect pubescence, on 6, towards the lateral
teeth, becoming longer and directed mesad.
Integument black with extensive pale areas, the ap-
pendages and metasoma predominantly pale; moder-
ately strongly, moderately densely to subreticulately
punctate, microsculpture not evident at 37,53.
Distribution. — N.E. Africa.
Host. — Not known.
Type species. — Aethammobates prionogaster, sp. n.
De nominis. — Gr. i En +
Ammobates, nom. propr. Hymenoptera
Aethammobates is separated from other nomadines
by a variety of autapomorphies, but for purposes of
recognition its general habitus and the modifications
of the pronotum and first metasomal tergum are im-
mediately diagnostic (see Discussion).
Aethammobates prionogaster Sp. n.
(figs. 1-4)
Type material. — Holotype d : ‘Coll. A. Mochi / 27.V.37
/ Geb. Asfar / Egitto’ (in coll. Baker )
Description
Male. — Structural characters: See generic descrip-
156
tion. POL = OOL. Hamuli 8. Length 7.5 mm, forewing
4.75 mm.
Vestiture. The face, dorsal surface of pronotum,
peripheries of mesoscutum and scutellum, mesepi-
sterna, and outer ventral surfaces of posterior coxae,
all more or less densely clothed with white squami-
form or subsquamiform pubescence, that on the pro-
notum apparently particularly dense (matted in type);
elsewhere on head and mesosoma sparser, the individ-
ual hairs becoming longer and less squamiform.
Terga with broader, white, medially emarginate, lat-
erally expanded, basal fasciae and narrow, entire, api-
cal fasciae, filling the depressed marginal areas, of
squamiform pubescence; tergum 5 nearly completely
covered. Disc of pygidial plate with similar pubes-
cence. Sterna 2-4 with weak submarginal fasciae, ster-
na 5 and 6 medially with fine, dense, erect pubes-
cence (as noted in generic description - possibly
homologous with pre-genital brush in Nomada).
Integument of head and mesosoma black with ex-
tensive pale areas. Clypeus, anterior extremities of
paraocular areas, malar areas, labrum, small maculae
adjacent to summits of eyes, larger maculae on genal
areas adjacent to mandibles, extending narrowly up-
wards almost to summits of eyes, labrum and all head
appendages (except the mandibles apically) castane-
ous, the flagellum darker. Lateral lobes of pronotum,
axillae, tegulae, sclerites of wing bases, wing veins (the
costa darker), and legs (the coxae basally darker) cas-
taneous. Metasoma castaneous, terga 2-5 discally pro-
gressively, irregularly (possibly the result of post-mor-
tem decomposition) darker; sterna 2 and 3 discally
largely, 4 and 5 almost wholly, dark. Labrum smooth,
moderately strongly, irrergularly punctate; anterior
face of pronotum smooth, impunctate; mesoscutum
and scutellum smooth, moderately densely, simply
punctate; terga smooth, medially densely, laterally re-
ticulately, punctate, the punctation coarsest on ter-
gum 2, on tergum 1 about equal to that of mesoscu-
tum, on terga 3-5 progressively finer. Margins of
pygidial plate reflexed. 1 J
Derivatio nominis. — Gr. TPLOV — WdNs / WTOS,
serrated, + gaster.
Discussion
Both Alexander (1990) and Roig-Alsina (1991)
have recently published cladistic analyses of nomadi-
ne bees. Roig-Alsina’s analysis was based primarily
(15 out of 22 characters) on characters exclusive to
the female, and is consequently of limited use in the
present context. Alexander’s analyses were based on
(1) larval characters only, (2) on adult characters of
those genera where larvae were known, (3) on both
larval and adult characters of these genera, and (4) on
the adult characters of all genera known to him. It is
this last analysis, (4), that is relevant to the placement
Figs. 1-4. Aethammobates prionogaster sp.n.
157
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Table 1 Data matrix for characters as listed by Alexander, Appendix 6; codings in accordance with Alexander, Appendices 3
and 5.
Characters
Holcopasites
1
1
Schmiedeknechtia 1
Aethammobates 0
SU
nN
Schmiedeknechtia
1
Holocopasites 0
0
Aethammobates I
© © © =
Schmiedeknechtia
2
Holocopasites 1
1
Aethammobates I
— eS ©
Sy) Inn)
Notes on matrix
a 6 7 8 9 10 11
1 0 0 1 0 0 0
“+ 0 0 0 0 0 0
0 0 = 0 1 0 2
16 17 b c 18 d 19
0 1 1 0 Ik 1 0
0 1 1 0 Ik 1 0
0 0° 0 1 1 1 1
g 22 h 23 24 29 32
0 0 2 1 0 1 0
0 0 D 1 0 1 0
0 0 2 if 0 0 07
(1) character a: Antennal sockets below mid-level of eyes, but not far below; no code. — (2) character 7: ‘Length/width of first
flagellomere’ = 3: no code. — (3) character 17: No ‘patches of appressed squamiform setae’: Alexander presumably refers to
discrete metasomal maculations of Holocopasites and Schmiedeknechtia. — (4) character 18: Incorrectly coded ‘0° by Alexander:
vein lst r-m of forewing is absent (two submarginal cells). — (5) character g: ‘arising from’ in Alexander’s Appendix 5 presum-
ably means ‘rising from’ (i.e, in lateral aspects bends upward from ventral plane of mesosoma immediately before intermedi-
ate coxae). — (6) character 23: a weak, short, dorso-basal carina present on posterior coxa. — (7) character 32: Coded ‘0’ since
apex narrowly rounded, subacuminate, but galea is more than weakly sclerotized.
of Aethammobates. Alexander's data matrix (his
Appendix 6) employed 46 characters, of which six ap-
plied solely to the female, leaving 40 in which
Aethammobates could potentially be compared. In
practice, since the holotype and only known speci-
men of Aethammobates has not been dissected (cf.
Roig-Alsina’s comment, 1991: 25, on the pre-epis-
ternal sulcus in Rhopalolemma), comparison has been
limited to 33 characters. These have been covered in
the description given above, and are now given in co-
ded form, following Alexander’s Appendices 3 and 5,
as an extension of his data matrix, in Table 1. This ta-
ble reproduces Alexander’s codings, but substitutes
observed values for six characters marked with a quae-
rein his matrix.
It should be noted that Alexander’s codings, here
and elsewhere, cannot always be accepted because he
did not see certain described and undescribed taxa
that would have necessitated different coding or the
adoption of additional codes; however, a complete re-
vision of his data matrix, which might well result in
significant changes in his cladogram, is beyond the
scope of the present paper. Important characters that
should be taken into account in future work on the
phylogeny and classification of the Nomadinae in-
clude the presence of facial foveae in some Nomada (a
group of, chiefly, Panurgus parasites); the develop-
ment of an anal truncation, with associated modifica-
tions of the apical sterna, among the pasitines
(Bischoff 1923: 585, Analstutz); and modifications of
158
the antenna in such genera as Morgania and
Pasitomachthes, where the scape and pedicel may be
coalescent, forming a single functional unit (Baker
1971: 7, footnote).
Inspection of the extended matrix confirms a prio-
ri impressions that Aethammobates shares a preponde-
rance of characters with Holcopasites and Schmie-
deknechtia and must be placed in the Holcopasitini.
While obviously most nearly related to Holcopasites
[Nearctic, revised by Linsley (1943, as Neopasites) and
by Hurd and Linsley (1972)] and Schmiedeknechtia
[western palaearctic, revised by Popov (1933)], and
sharing such derived characters as the lateral process-
es of the metanotum and certain venational details,
Aethammobates is not close to either. It differs from
both in, among other characters:
1. The dorso-ventrally compressed form of mesoso-
ma and metasoma.
2. The mid-dorsally exposed pronotum with sharply
differentiated dorsal, densely pubescent and anter-
ior, glabrous surfaces. (In lateral aspect, the prono-
tum resembles that of Pae (Sphecoidea) as illus-
trated by Bohart & Menke (1976: 15, fig. 3A),] In
Holcopasites and Schmiedeknechtia the pronotum
is, as in most other bees, recessed beneath the me-
soscutum with only its lateral lobes and, when the
head is deflexed, the collum conspicuous in dorsal
aspect.
3. The V-shaped interantennal carina.
4. The strongly expanded intermediate femora.
5. The pronounced basal truncation of the mesoso-
ma.
6. The form of the first metasomal tergum, which is
broadly, deeply emarginate, with prominent ante-
ro-lateral angles (see fig. 3) and the anterior and
dorsal surfaces sharply, rectangularly separated.
7. The presence of basal and marginal tergal fasciae
and the absence of of discrete spot- or bar-like
markings.
Of these, characters 1-6 appear as autapomorphies.
Aethammobates differs further, from male
Holcopasites, in the 13-segmented antennae, and from
Schmiedeknechtia in the form of the pygidial plate,
which is not narrowed basad; in not having the inner
orbits convergent below; in not having the anterior
mandibular articulations in contact with the eyes; and
in the venation: in Schmiedeknechtia either both 1 m-
cu and 2 m-cu are received by second Rs, or
(Cyrtopasites, a single species with convex mesoscu-
tum and fine and sparse punctation) 1 m-cu is inter-
stitial with or slightly precedes 1 r-m.
Biology
It is interesting to speculate on the possible host of
Aethammobates. Known hosts of Holcopasites and
Schmiedeknechtia are panurgines. Camptopoeum and
Meliturgula are possible candidates, but the former.
which is normally abundant where it does occur, and
which appears not to have been recorded from the
Cairo area (it is not represented in recent, extensive
Egyptian collections), has species of Parammobatodes
as its regular parasites. The somewhat flattened shape
of Aethammobates suggests some correlation with
Baker: A new genus of nomadine bees
habits or host. Both sexes of Meliturgula have a cons-
picuously broad or flattened metasoma: regional spe-
cies are few in number and rare in collections; not-
hing is known of their biology.
REFERENCES
Alexander, B., 1990. A cladistic analysis of the nomadine
bees (Hymenoptera: Apoidea). — Systematic Entomology
1522151572
Baker, D. B., 1971. A new Pasitomachthes from Rhodesia
(Hymenoptera, Apoidea). — Novos Taxa entomológicos,
no. 98: 1-8.
Bischoff, H., 1923. Zur Kenntnis
Schmarotzerbienen. — Deutsche
Zeitschrift 1923: 585-603.
Bohart, R. M. & A. S. Menke, 1976. Sphecid wasps of the
world: a generic revision: ix + 695 pp. — University of
California Press, Berkeley.
Hurd, P. D. & E. G. Linsley, 1972. Parasitic bees of the ge-
nus Holcopasites Ashmead (Hymenoptera: Apoidea). —
Smithsonian Contributions to Zoology, no. 114: 1-41.
Linsley, E. G., 1943. A revision of the genus Neopasites
(Hymenoptera: Nomadidae). — Transactions of the
American Entomological Society 69:119-140.
Popov, V. B., 1933. Notes on the parasitic bees allied to the
genus Biastes Panz. (Hymenoptera, Nomadidae). —
Trudy zoologicheskogo Instituta Akademy Nauk SSSR 2:
51-75.
Roig-Alsina, A., 1991. Cladistic analysis of the Nomadinae
s.str. with description of a new genus (Hymenoptera:
Anthophoridae). — Journal of the Kansas Entomological
Society 64: 23-37.
afrikanischer
entomologische
Received: 7 April 1993
Accepted: 30 December 1993
159
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
160
DEB OER
Institute for Systematics and Population Biology (Zoölogisch Museum), University of Amsterdam
ROURSSLECIESFADDEPD MO NE BAF KURIA NASCITA
GROUP, WITH NOTES ON TAXONOMY AND
BIOGEOGRAPHY (HOMOPTERA, TIBICINIDAE)
Boer, A. J. de, 1994. Four species added to the Baeturia nasuta group, with notes on taxonomy
and biogeography (Homoptera, Tibicinidae). — Tijdschrift voor Entomologie 137: 161-172,
figs. 1-39. [ISSN 0040-7496]. Published 15 December 1994.
A new concept is proposed for the nasuta group of the cicada genus Baeturia Stal, 1866. Four
species are added to this group, bringing the total number of its species to twelve. One of these
species (B. guttulipennis Blöte, 1960) is redescribed and three (B. gibberosa, B. splendida, and B.
retracta) are described as new. Implications for the biogeographic pattern of the group are, that
the nasuta group can no longer be seen as a typical central New Guinea group; most of the spe-
cies can apparently be found in northern New Guinea.
AJ. de Boer, Institute for Systematics and Population Biology (Zoölogisch Museum),
University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands
Key words. — Baeturia, nasuta group, taxonomy, biogeography, New Guinea.
The nasuta group was recognized as a monophylet-
ic group consisting of eight New Guinean species of
the genus Baeturia Stal (de Boer 1982). The mono-
phyly of that group is based on two presumed apo-
morphies: a narrow and angularly swollen postclype-
us and four sclerotized ridges on the tymbals. B.
guttulipennis Blöte was not included in the nasuta
group, since it does not share these apomorphies.
However, now that most of the remaining species of
Baeturia have been revised and placed in monophy-
letic species groups, it appears that B. guttulipennis is
more closely related to the nasuta group than to any
other species group. It is proposed here to change the
concept of the nasuta group and to include B. guttu-
lipennis in that group. Furthermore, three new species
can be added to the nasuta group, these species do
share the original apomorphies of that group.
Species of the nasuta group, as previously defined,
were mainly distributed in the central mountain
ranges of New Guinea. Its distribution pattern has
been compared to that of the cicada genus
Cosmopsaltria Stal, which also centres in the montane
parts of central New Guinea (de Boer 1982; Duffels
1986; Duffels & de Boer 1990) and it was suggested
that the similarities between these patterns might find
a common cause in the paleogeology of New Guinea.
The present additions to the nasuta group greatly al-
ter its general distribution pattern and necessitate a
reconsideration of its paleogeographic history.
MATERIAL AND METHODS
The material examined for this study is deposited
in the following collections:
AMNH: American Museum of Natural History,
New York; AMs: Australian Museum, Sydney; BMNH:
Natural History Museum (formerly: British Museum
(Natural History)), London; BPBM: Bernice P. Bishop
Museum, Honolulu; Moul: Personal collection Mr
M.S. Moulds, Sydney; RMNH: Nationaal Natuur his-
torisch Museum (formerly: Rijksmuseum van
Natuurlijke Historie), Leiden; SMN: Staatliches
Museum für Naturkunde, Stuttgart; ZMA: Institute
for Systematics and Population Biology (Zoölogisch
Museum), Amsterdam.
The following geographical sources have been
used: Atlas van tropisch Nederland (1938), The
Times Atlas of the World (1968) and the ‘List of New
Guinea localities’ published by the Bishop Museum
(1966).
After overnight softening, male genitalia were ex-
amined by pulling out the pygofer with a sharp nee-
dle inserted between pygofer and 8th abdominal seg-
ment. The aedeagus was pulled out at the same time,
by inserting the needle between the claspers.
Measurements are based on all available specimens.
PHYLOGENY
Baeturia belongs to a larger group of genera, the
161
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
‘Baeturia and related genera complex’, as defined ear-
lier (de Boer 1990). A phylogenetic analysis of this ge-
nus complex, with the oriental Prasiini as defined by
De Jong (1985) as outgroup, is in preparation. The
ensuing phylogenetic discussions are based on the
preliminary results of that analysis.
The monophyly of the nasuta group
The monophyly of the nasuta group is based on
three presumed apomorphies. Males of the nasuta
group are easily identified by the almost transparent
lateral sides of the abdomen, usually restricted to seg-
ments 3-7. Only B. marmorata does not show this
character. A similar transparency of the male abdo-
men was found in a monophyletic group of three spe-
cies of the genus Gymnotympana (unpublished), but
not in other related taxa. A laterally transparent abdo-
men is regarded as apomorphous for the nasuta group
(1 in fig. 1b) and the similar transparency in three
Gymnotympana species must be explained by paralle-
lism.
A strongly curved aedeagus, with lateral lobes at its
basal curve, is regarded apomorphous for the genus
Baeturia. The aedeagus of the species of the nasuta
nasuta group
5 viridis group
4 conviva group
6 guttulinervis group
1 2 3 bloetei group
exhausta group
1a loriae group
laminifer
9 retracta
8 bipunctata
gibberosa
6 7 arabuensis
mamillata
marmorata
AUS parva
splendida
0283 intermedia
nasuta
ib guttulipennis
Fig. 1. Cladograms. — a. Cladogram of Baeturia, numbers
refer to characters discussed in the section on the phyloge-
netic position of the nasuta group; b. Cladogram of the
Baeturia nasuta group, numbers refer to characters discussed
in the sections on the monophyly and ingroup phylogeny of
the „asuta group.
162
group differs from that of other species by a longer
and more slender apical part, distally of the lateral
lobes (fig. 11), which is regarded apomorphous (2 in
fig. 1b). A very similarly elongate, but stouter, aedea-
gus was found in B. loriae and B. pigrami of the lori-
ae group (de Boer 1994a). The elongation of the ae-
deagus in these two groups cannot be regarded as
homologous in the most parsimonious solution.
The male pygofer is characterized by a strongly
bent and dorsally flattened caudodorsal beak. This
shape of the caudodorsal beak is shared by all species
and regarded apomorphous for the nasuta group (3 in
fig. 1b). In other species, which have a similarly
strongly bent beak, the beak is convexly curved dor-
sally (comp. the viridis group, de Boer 1992).
The phylogenetic position of the nasuta group
Baeturia is a monophyletic genus, for which a fron-
tally rounded and protruding postclypeus, a triangle-
shaped medial thorn on the fore femur, and lobate la-
teral crests on the aedeagus are the presumed
apomorphies (1, 2, 3 in fig. la). Baeturia can be sub-
divided into seven monophyletic species groups: the
bloetei group (de Boer 1989), the conviva group (de
Boer 1986), the exhausta group (de Boer 1994b) the
guttulinervis group (de Boer 1994c), the loriae group
(de Boer 1994a), the viridis group (de Boer 1992),
and the nasuta group (de Boer 1982).
The nasuta group is presumed to form a monophy-
letic group with the conviva, guttulinervis, and viridis
groups. The large, often conically-shaped, distinctly
posteriorly projecting protuberance on the lateral
lobe of pygofer is regarded synapomorphous for these
four species groups together (4 in fig. la).
Furthermore, these four groups share a very stout and
strongly bent caudodorsal beak. The occurrence of a
similarly bent caudodorsal beak in several Melanesian
species of the bloetei group must be explained by par-
allel development.
The nasuta and viridis groups are sister groups,
sharing an angularly bent caudodorsal beak as sup-
posed synapomorphy (5 in fig. la). In the species of
these two groups the dorsal margin of the pygofer is
not continuously rounded with the dorsal margin of
the caudodorsal beak. Furthermore, these species
share a generally truncate, but sometimes bluntly
rounded or bicuspidate, caudodorsal beak. Species of
other groups of Baeturia generally have a more nar-
rowly rounded or pointed beak, while a truncate beak
only sporadically occurs. The conviva and guttuliner-
vis groups are sister groups based on a shared apical
swelling of the clasper (this swelling is almost globu-
lar in the guttulinervis group) and a dorsally strongly
rounded pygofer (6 in fig. la), but the relative posi-
tion of the remaining species groups of Baeturia is not
clear.
Ingroup phylogeny
The species of the nasuta group are very similar in
general appearance. The species differ mainly in the
shape of the claspers and some clasper characters indi-
cate phylogenetic relationships. Most other differen-
ces observed are either species specific characters, or
characters that also occur outside the nasuta group. It
is not possible to construct a fully resolved and un-
ambiguous cladogram for the group, based on the li-
mited number of characters now available.
Nevertheless, some remarks on ingroup phylogeny
can be made (see fig. 1b).
B. guttulipennis presumably forms the sister taxon
of all other species of the nasuta group; these other
species share four tymbal ridges (three in B. parva)
and a more angularly swollen postclypeus, which are
the original apomorphies for the asuta group (4, 5 in
fig. 1b). B. guttulipennis and B. nasuta however, share
similar brown markings on the tegmina and an api-
cally rounded male caudodorsal beak, and could be
sister species. B. nasuta and B. intermedia share an al-
most identical clasper and are presumably either sister
species or part of a paraphyletic subgroup.
B. arabuensis, B. bipunctata, B. laminifer, B. ma-
millata, B. retracta and B. gibberosa share a distinct
dorsal protrusion on the clasper as presumed synapo-
morphy (6 in fig. 1b). That B. nasuta and B. interme-
dia should also be included in this group, as was sug-
gested before (de Boer 1982) seems unlikely, since the
weakly developed dorsal crest of the claspers of B. na-
suta and B. intermedia is very similar to that found in
many other Baeturia species. The remaining species
of the nasuta group share a dorsally smoothly round-
ed clasper.
De BOER: The Baeturia nasuta group
B. arabuensis and B. mamillata share a posteriorly
directed and almost finger-shaped protrusion, and are
possibly sister species. B. bipunctata, B. gibberosa, B.
laminifer and B. retracta have a much broader dorsal
protrusion on the clasper, which is regarded as synap-
omorphous (8 in fig. 1b). B. laminifer and B. retracta
share a laminiform and square-shaped dorsal protru-
sion on the clasper as presumed synapomorphy (9 in
fig. 1b). B. bipunctata and B. gibberosa have a more
triangular, hump-shaped, protrusion; these are either
sister species or part of a paraphyletic subgroup.
Females of most of the species that have a dorsal
protrusion on the clasper, as far as they could be iden-
tified, generally have a truncate caudodorsal beak,
which character is unique for these species and pre-
sumably apomorphous at that level (7 in fig. 1b).
Only the females that presumably belong to B. ara-
buensis and the females of B. gibberosa have a pointed
beak as those in B. guttulipennis, B. nasuta, and B.
parva. The female of B. splendida has a bluntly round-
ed, almost truncate, caudodorsal beak.
BIOGEOGRAPHY
The nasuta group is distributed in New Guinea
and some adjacent islands, some species reach to the
Bismarck Archipelago or the Admiralty Islands.
Study of the New Guinean cicadas indicates that
the various genera or monophyletic species groups
show different patterns of distribution. The genera
seem concentrated with most, and often endemic,
species in different parts of the island. These parts of
New Guinea, thus recognized as areas of endemism
(Duffels & de Boer 1990), largely coincide with frag-
Tren SSE SZ I 1 = 5 | | |
| | Fa) Fig. 2. Distribution of
B guttulipennis Baeturia gibberosa, B. gut-
@ retracta tulipennis, B. retracta, and
à ALERTE ms B. splendida.
i Y gibberosa [|
b> È ESS
( SEZ = ED ||
1 wu.
A :
à 5
È i
0. Da 5
Lai 2 È io 4
IS)
Us ee
©
LA DS > : ® x 8°
SS Ge
S a oc
Olle © u, : OP Es
Sa i 5 = 4
ie nu, See Pac
1 I I
136° 140° 144° 148° 152°
163
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
ments of a historic island arc, that collided at different
times and in various places with the northern craton
of the Australian continent, to form present day New
Guinea (for geological reviews see Daly et al. 1991;
Rangin et al. 1990a, 1990b; Pigram & Davies 1987).
It is supposed that the various genera of New
Guinean cicadas evolved on isolated island arc frag-
ments previous to their collision with the Australian
continent (de Boer 1994c).
The nasuta group was regarded as a typical central
New Guinea group (de Boer 1982) and, as such, has
been compared to the genus Cosmopsaltria Stal
(Duffels 1986; Duffels & de Boer 1990).
Cosmopsaltria and the B. nasuta group both appeared
to be concentrated in the central mountain ranges of
New Guinea. The distributions of B. arabuensis, B.
mamillata and B. nasuta, from the Wissel Lakes to
well into the Papuan Peninsula, are indeed very simi-
lar to that of many species of Cosmopsaltria (Duffels
1983). A similar historical explanation for the distri-
bution patterns found in these two groups seemed re-
quired. Such an explanation can be found in suppos-
ing that both Cosmopsaltria and the nasuta group
evolved on the Sepic Arc terrane as defined by Pigram
& Davies (1987), which is the first of the island arc
fragments that collided with the Australian continent,
and which developed into the central mountain rang-
es of New Guinea. However, the phylogenetic level of
the groups (Cosmopsaltria as sister group of
Diceropyga, Rhadinopyga, Aceropyga, and Moana
(Duffels 1986, 1993), and the nasuta group as a sub-
group of Baeturia) possibly indicates a considerable
age difference, which contradicts to a contemporal
evolution on the Sepic Arc fragment.
With the four species here added to the nasuta
group, this group as a whole no longer has a typical
central New Guinea distribution. In fact, the number
of species that occurs in northern New Guinea about
equals that of central New Guinea. Only four species
(B. arabuensis, B. intermedia, B. mamillata, and B.
nasuta) can be said to have a central mountain range
distribution, though two (B. mamillata, and B. nasu-
ta) also occur in the Papuan Peninsula. The distribu-
tions of B. laminifer and B. retracta are very similar;
they are recorded from southern New Guinea (un-
published data for laminifer), the most eastern part of
northern New Guinea, and the Papuan Peninsula. B.
laminifer is recently recorded from New Britain (un-
published) and B. retracta is recorded from the
Admiralty Islands. These two species are absent from
the western parts of the central mountain ranges. B.
gibberosa is possibly restricted to the Papuan
Peninsula and Normanby Island. B. splendida and,
possibly, B. guttulipennis are endemic to northern
New Guinea (it is not certain that the female allotted
to the latter species really belongs to B. guttulipennis).
164
Also B. parva is distributed in northern New Guinea,
but its distribution extends to the Vogelkop
Peninsula, Misool, Biak, the Bismarck Archipelago,
and Admiralty Islands. The two remaining species (B.
bipunctata and B. marmorata), both from Araucaria
camp and Rattan camp on the northern slopes of the
central mountains of western New Guinea, should
presumably be regarded more as northern, than as
central New Guinean: several other species groups
with a predominantly northern New Guinea distri-
bution (e.g. the viridis group and the guttulinervis
group) also have species in that area.
The distribution pattern of the nasuta group in its
present concept is more in agreement with that of
other groups of Baeturia, most of which seem to con-
centrate in northern New Guinea, while similarities
with Cosmopsaltria have become less obvious and a
paleobiogeographical comparison with that genus
seems no longer relevant.
‘TAXONOMY
Description of the nasuta group
Species of the nasuta group are all very small sized.
The specimens are shorter than 2.5 cm and generally
do not exceed 2.0 cm. Males are easily recognized by
their laterally transparent abdomen (this character is
absent in B. marmorata). Body generally yellowish
brown or reddish brown coloured and covered with
many irregularly shaped brown speckles. Females ge-
nerally darker coloured and more densely speckled
(B. splendida unspeckled with greenish head and tho-
rax).
Head (fig. 3a): Narrow, 2.3-2.8 x as wide as long.
Vertex narrower than in most other Baeturia species,
ocelli close together. Distance between lateral ocelli
less than 2 x as wide as, and sometimes narrower than,
frontal ocellus, and generally about as long (0.7-1.2 x)
as distance between lateral ocellus and eye. Vertex
1.6-2.5 x as wide as postclypeus. Postclypeus in dorsal
view narrow (1.1-2.2 x as wide as long) and triangu-
larly protruding beyond vertex lobes, though with
rounded anterior margin. Postclypeus distinctly swol-
len ventrally, its anterior margin (lateral view) often
angularly bent (fig. 3b).
Legs: Fore femur (Fig. 6) with row of three erect
and sharply pointed spines, diminishing in length to-
wards tibia. The most distal spine often bicuspidate,
forming a very small 4th spine. Most proximal spine
slightly shorter than distance to middle spine. Middle
spine triangular, very broad at its base.
Tegmina and wings: Hyaline, though tegmina
sometimes with darkly suffused spots along veins.
Tegmina with 8, wings with 6 apical areas. Hyaline
border along hind margin of tegmen distinct, though
narrow. This border slightly broader in wing. Veins
ochraceous, sometimes tinged with red, and set with
many long setae.
Tymbals: With 3-5, generally 4, parallel sclerotized
ridges from dorsal to ventral tymbal margin. Often
traces of red between dorsal parts of tymbal ridges.
Opercula: Male operculum fairly small, its distal
part not covering tymbal cavity in ventral view and
nearly always shorter than meracanthus, generally not
reaching to anterior margin of 2nd abdominal seg-
ment. Operculum in ventral view reaching to under,
or medially of, meracanthus. Female operculum
smaller than that of male, sickle-shaped and erect.
Abdomen: Male abdomen conspicuous by almost
transparent sides, its dorsal and distal parts darkened
and brown speckled, ventral parts unspeckled.
Lateroventral row of dark spots generally very dis-
tinct. Hind margins of abdominal segments often
reddish. First and second sternites adjacent. First ter-
gite very short and often partly hidden under metano-
tum (in B. arabuensis distinctly longer than in other
species). Second tergite medially with almost straight
anterior margin. Auditory capules well developed,
globularly swollen. Female abdomen more robust
than that of male, darker coloured, and more densely
speckled. Ovipositor sheaths sometimes reaching be-
yond apex of caudodorsal beak. Female caudodorsal
beak often truncate at apex.
Male genitalia: Caudodorsal beak strongly bent,
making an almost right angle with dorsal margin of
pygofer. Dorsal margin of pygofer not continuously
rounded into dorsal margin of beak, and weakly con-
cave near apex of beak. Caudodorsal beak very broad
and straight, dorsally flattened, and with bluntly
rounded or truncate, often weakly concave, apex.
Lateral lobe of pygofer with strongly swollen, often
conically shaped, protuberance, projecting posterior-
ly beyond margin of pygofer. Claspers fairly long,
parallel, and generally directed posteriad. Apical part
of clasper hardly bent down, with large and sharply
edged ventral hollow. Aedeagus S-curved with lateral
lobes at basal curve. Apical part of aedeagus extreme-
ly long and slender (compared to other species of
Baeturia). Aedeagal pore narrow and oval.
Baeturia guttulipennis Blöte
(figs. 2-13)
Baeturia guttulipennis Blote, 1960: 69, fig. 15; de Boer,
1994c.
Material examined: Bewani R. territ., 1200 m, 1939, W.
Stüber, d holotype, RMNH; Bivak 39A, Star Range, 1150
m, 6.vii.1959, 12 Baeturia guttulipennis ? det. H.C. Blôte,
RMNH.
B. guttulipennis is easily separated from other spe-
DE BOER: The Baeturia nasuta group
Figs. 3-6. Baeturia guttulipennis Blote, 1960. — 3a, head in
dorsal view, holotype; 3b, head in lateral view, holotype; 4,
female caudodorsal beak in dorsal view; 5, female genital
segment in lateral view; 6, fore femur, holotype.
cies of the nasuta group by the dark spots along the
veins of tegmina and by its five tymbal ridges. Similar
dark spots in tegmina were found in several other spe-
cies of Baeturia, e.g. B. guttulinervis Blöte (de Boer
1994c), B. pigrami De Boer and B. silveri De Boer (de
Boer 1994a), and, though less distinct, in B. nasuta
Blôte (de Boer 1982).
Description
Body of male light ochraceous brown and weakly
speckled with brown. Female darker, red-brown and
more densely speckled. Female head and thorax larger
than in male. Tegmina of males 1.2 x as long as body
length, of females 1.5 x. Male abdomen 1.3 x as long
as head and thorax, of females 1.1 x.
Head (fig. 3a): Ochraceous, brown-stained mainly
on vertex lobes. Postclypeus bluntly rounded anteri-
orly, weakly protruding in male, more strongly in fe-
male. Postclypeus 1.4-2.0 x as wide as long.
Postclypeus in lateral view (fig. 3b) distinctly swollen,
anterior margin forming an almost right angle at 1/2
its length, and concave to anteclypeus.
Thorax: Pronotum ochraceous with dark speckling
predominantly in medial band and on medial parts of
165
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
18 19 du fo 20 21
Figs. 7-13. Baeturia guttulipennis Blöte, 1960: 7, pygofer in lateral view, holotype; 8, clasper, holotype; 9, male operculum,
holotype; 10, male caudodorsal beak in dorsal view, holotype; 11, aedeagus from behind, holotype; 12, aedeagus in lateral
view, holotype; 13, female operculum. 14-22. Baeturia splendida sp. n.: 14, pygofer in lateral view; 15, male caudodorsal
beak in dorsal view; 16, clasper; 17, male operculum; 18, head in lateral view; 19, female operculum; 20, female caudodorsal
beak in dorsal view; 21, aedeagus in lateral view; 22, aedeagus from behind. Lettering: bp = basal part of operculum; cb = cau-
dodorsal beak; d = distal margin of operculum; di = = distal margin of pygofer; dmc = distomedial corner of operculum; do =
dorsal margin of pygofer; dp = distal part of operculum; he = clasper heel; ho = clasper hollow; | = lateral margin of opercu-
lum; m = medial margin of operculum; p = protuberance on lateral lobe of pygofer; ve = ventral margin of pygofer.
166
pronotal lobes, unspeckled in and along its fissures.
Mesonotum greenish brown, weakly brown speckled
in male, but densely speckled in female. Male with
two brown spots in front of cruciform elevation.
Legs: Ochraceous, often reddish towards tarsi, and
brown speckled. Fore femur (Fig. 6) with row of three
or four sharply pointed and erect spines.
Tegmina and wings: Tegmina with dark brown
markings concentrated along veins. Wings hyaline.
Veins ochraceous brown, densely set with setae.
Tymbal organ: Five darkly sclerotized parallel ridg-
es spanning the tymbal from dorsal to ventral margin.
Four intercalary ridges form a lateral band across tym-
bal.
Opercula: Male operculum (fig. 9) very small and
not covering tymbal cavity. Distal part of operculum
erect and reaching to about 2/3 the length of mera-
canthus, but not reaching to abdomen. Lateral mar-
gin of distal part short, concavely bent into crest
around distolateral corner of basal part, and convexly
bent into long and almost straight distal margin.
Distomedial corner rounded, medial margin straight.
Female operculum (fig. 13) almost as large as that of
male, broadly sickle-shaped and curved to abdomen,
with almost continuously rounded distal margin.
Abdomen: Male abdomen weakly inflated, yellow
brown and brown speckled, though ventrally un-
speckled. Tergites 3-6 almost transparent laterally. A
distinct lateroventral row of brown spots on tergites
3-7. First tergite very short and medially partly hid-
den under metanotum. Female abdomen more ro-
bust than that of male, dark reddish brown, and
densely speckled. Lateroventral spots on tergites 3-7
almost lost in speckling. Ovipositor sheaths not
reaching to apex of caudodorsal beak (fig. 5). Female
caudodorsal beak in dorsal view (fig. 4) sharply point-
ed at apex.
Male genitalia: Pygofer in lateral view as in fig. 7.
Dorsal margin weakly convex, strongly bent into al-
most straight caudodorsal beak. Distal margin con-
cave, gradually bent into straight margin of beak.
Protuberance on lateral lobe of pygofer rounded and
conically projecting posteriad. Pygofer lobe forming
an angular corner just above this protuberance.
Ventral margin almost straight, but angularly bent
just under lateral protuberance. Ventral margins con-
verge to sharp angle at base of pygofer opening.
Caudodorsal beak in dorsal view (fig. 7) rounded at
apex. Clasper (fig. 8) almost straight and directed
posteriad, with long and slender, dorsally smoothly
rounded, apical part. Basal part of clasper with weak-
ly developed clasper heel. Clasper hollow fairly long.
Aedeagus (fig. 12) with very long and slender apical
part and well developed lateral lobes. Aedeagus from
behind (fig. 11) very slender, aedeagal pore narrow,
oval shaped and truncate at apex.
De BOER: The Baeturia nasuta group
Measurements: Body length d: 18.6 mm, 9: 18.2
mm; tegmen length d: 22.9 mm, 9: 28.0 mm; head
length d: 1.4 mm, 9: 1.8 mm; pronotum length d:
2.4 mm, 9: 2.8 mm; mesonotum length d : 3.8 mm,
?: 4.8 mm; head width d: 3.7 mm, 9: 4.3 mm;
width of pronotal collar d: 5.3 mm, 9: 6.5 mm.
Distribution (fig. 2). — The male holotype of B.
guttulipennis comes from Bewani in northeastern
Irian Jaya; a female that possibly belongs to the spe-
cies comes from the Star Range in central eastern
Irian Jaya.
Baeturia splendida sp. n.
(figs. 2, 14-22)
Holotype male: “NEW GUINEA NE / Torricelli Mts. /
Mokai Vill. 750 m / xii-16-31-1958° (print); “W.W. Brandt
/ Collector / Bishop’ [print], BPBM. — Paratypes: same data as
holotype 52, BPBM; same data but 8-15.xii.1958, 32; 1-
23.1.1959, 19, all BPBM.
Only one male and several females are known of
this species. B. splendida is easily recognized by its
greenish colour, with a pattern of brown markings.
Due to this colour pattern the females too are easily
identified. The colour pattern is unique within the
nasuta group.
Description
Head and thorax olive green, abdomen yellowish
brown. Tegmina of male 1.2 x as long as body length,
of females 1.3-1.4 x. Male abdomen 1.2 x as long as
head and thorax, of females 0.9-1.1 x.
Head: Olive-green. Vertex lobe with dark brown
spot between eye and lateral ocellus. Postclypeus dis-
tinctly protruding, bluntly rounded anteriorly, and
1.1-1.8 x as wide as long. Postclypeus in lateral view
(fig. 18) distinctly and angularly swollen, anteriorly
almost globularly inflated.
Thorax: Pronotum greenish and unmarked.
Mesonotum green with two large semi-oval parame-
dian brown spots at pronotal margin, reaching to
about half-length of mesonotum. No dark spots in
front of cruciform elevation.
Legs: Greenish, yellow-brown towards tarsi. Fore
femur with row of four sharply pointed and erect
spines.
Tegmina and wings: Hyaline. Veins greenish, and
set with setae.
Tymbal organ: Four weakly sclerotized parallel
ridges spanning the tymbal from dorsal to ventral
margin. Three intercalary ridges form a lateral band
across tymbal.
Opercula: Male operculum (fig. 17) very small and
not covering tymbal cavity. Distal part of operculum
erect and reaching to about half-length of meracan-
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
168
thus, not reaching to abdomen. Lateral margin of dis-
tal part short, concavely bent into crest around disto-
lateral corner of basal part, and convexly bent into
long and almost straight distal margin. Distomedial
corner rounded, medial margin straight. Female op-
erculum (fig. 19) almost as large as that of male,
broadly sickle-shaped and erect, with almost continu-
ously rounded distal margin.
Abdomen: Male abdomen hardly inflated, yellow
brown, darker brown in distal part. Third and 4th
tergites almost transparent laterally. A distinct latero-
ventral brown spot on 3rd tergite and a slightly small-
er lateral spot on 8th tergite. First tergite very short
and medially partly hidden under metanotum.
Female abdomen more robust than that of male, light
brown, with a dark brown lateroventral spot on 3rd
tergite. Ovipositor sheaths not reaching to apex of
caudodorsal beak. Female caudodorsal beak in dorsal
view (fig. 20) weakly convex, almost truncate, at apex.
Male genitalia: Pygofer in lateral view as in fig. 14.
Dorsal margin weakly convex, angularly bent into al-
most straight caudodorsal beak. Distal margin con-
cave, almost continuously bent with margin of beak.
Protuberance on lateral lobe of pygofer bluntly
rounded and weakly projecting posteriad. Pygofer
lobe forming an angular corner just above this protu-
berance. Ventral margin angularly bent at about 1/3
its length and weakly concave towards base. Ventral
margins converge to sharp angle at base of pygofer
opening. Caudodorsal beak in dorsal view (fig. 15) al-
most truncate at apex, but ending in very small and
sharply pointed medial thorn. Clasper (fig. 17) curv-
ing upwards with long and slender, dorsally smoothly
rounded and recurving, apical part. Basal part of
clasper without clasper heel. Clasper hollow fairly
long. Aedeagus (fig. 21) with very long and slender
apical part. Basal part of aedeagus with small and an-
gular dorsolateral protuberances. Lateral lobes small,
hardly projecting in lateral view. Aedeagus from be-
hind (fig. 22) very slender. Aedeagal pore narrow,
oval-shaped and truncate at apex.
Measurements: Body length d: 16.5 mm, 9:
16.0-19.1 mm (x 18.0 mm + 0.9); tegmen length d:
20.6 mm, 9: 22.0-25.5 mm (x24.6 mm + 1.0); head
length d: 1.5 mm, 9: 1.6-1.9 mm (x 1.8 mm); pro-
notum length d: 2.3 mm, 9: 2.8-3.2 mm (x 2.9
mm); mesonotum length d : 4.0 mm, 9: 4.0-5.2 mm
(x 4.8 mm); head width ¢: 3.3 mm, 2: 4.0-4.8 mm
(x 4.6 mm); width of pronotal collar d : 5.1 mm, 2:
DE BOER: The Baeturia nasuta group
5.7-7.0 mm (x 6.6 mm).
Distribution (fig. 2). — B. splendida is only known
from the Torricelli mountains in northern Papua
New Guinea.
Etymology. — Splendidus (Latin) refers to the beau-
tiful colour pattern of this species.
Baeturia gibberosa sp. n.
(figs. 2, 23-31)
Holotype: ‘New Guinea SE / Popondetta / 25 m vi.66’ (-
print); ‘Shanahan- Lippert / Light Trap / Bishop Museum’
(print), d , BPBM. — Paratypes: NEW GUINEA: Popondetta, 60
m, 1-4.ix.1963, J. Sedlacek, 26, BPBM; Popondetta, Inbora
Plant’n, 23.ix.1963, P. Shanahan, 26, BPBM; NORMANBY:
Wakaiuna, Sewa Bay, 5-9.x1.1956, W.W. Brandt, 19,
BPBM; same data but 1-10.xii.1956, 19; 11-20.xii.1956,
19, both BPBM; Wamula, 400-500 m, 31.xii.1988, R. de
Keyzer, 3d, Moul.
B. gibberosa is easily recognized by the rectangular
dorsal corner of its clasper.
Description
Body ochraceous brown, sometimes tinged with
red, and densely speckled with brown. Tegmina of
males 1.1-1.2 x as long as body length, of females 1.3
x. Male abdomen 1.3-1.5 x as long as head and tho-
rax, of females 1.0 x.
Head: Ochraceous, brown speckled. Postclypeus
weakly protruding and bluntly rounded anteriorly,
1.7-2.2 x as wide as long. Postclypeus in lateral view
distinctly swollen with straight anterior margin, con-
cave towards anteclypeus.
Thorax: Pronotum ochraceous, brown speckled.
Mesonotum greenish brown, slightly brown speckled.
Two dark spots in front of cruciform elevation.
Legs: Ochraceous and weakly speckled. Fore femur
with row of four sharply pointed and erect spines.
Tegmina and wings: Hyaline. Veins ochraceous,
moderately set with setae.
Tymbal organ: Four weakly sclerotized parallel
ridges spanning the tymbal from dorsal to ventral
margin. Three intercalary ridges form a lateral band
across tymbal.
Opercula: Male operculum (fig. 30) small and on-
ly partly covering tymbal cavity. Distal part of oper-
culum oval shaped, slightly erect and reaching to
about 2/3 of length of meracanthus, in some speci-
mens reaching to anterior margin of 2nd abdominal
Figs. 23-31. Baeturia gibberosa sp. n. — 23, pygofer in lateral view; 24, pygofer from aslant; 25, clasper; 26, female caudodor-
sal beak in dorsal view; 27, aedeagus from behind; 28, aedeagus in lateral view; 29, male caudodorsal beak in dorsal view; 30,
male operculum; 31, female operculum
Figs. 32-39. Baeturia retracta sp. n. — 32, female caudodorsal beak in dorsal view; 33, pygofer in lateral view; 34, aedeagus
from behind; 35, aedeagus in lateral view; 36, pygofer from aslant; 37, male caudodorsal beak in dorsal view; 38, female op-
erculum; 39, male operculum.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
segment. Lateral margin of distal part fairly long and
directed mesiad, concavely bent into crest around dis-
tolateral corner of basal part, and forming an obtuse
angle with straight distal margin. Medial part of oper-
culum angularly rounded, extending medially of mer-
acanthus. Female operculum (fig. 31) short, sickle-
shaped and erect.
Abdomen: Male abdomen not inflated, ochraceous
brown and brown speckled, though ventrally un-
speckled. Segmental hind margins reddish. Tergites
3-6 almost transparent laterally. A distinct lateroven-
tral row of brown spots on tergites 3-8. First tergite
very short and medially partly hidden under metano-
tum. Female abdomen red-brown and densely speck-
led. Female caudodorsal beak (fig. 26) sharply point-
ed at apex.
Male genitalia: Pygofer in lateral view as in fig. 23.
Dorsal margin weakly concave, but convexly bent into
almost straight caudodorsal beak and concave near
apex of beak. Distal margin concave, and continuous-
ly bent into margin of beak. Protuberance on lateral
lobe of pygofer bluntly rounded and weakly projecting
posteriad. Pygofer lobe forming a small angular corner
just above this protuberance. Ventral margin almost
straight, but convexly bent just under lateral protuber-
ance. Ventral margins converge to sharp angle at base
of pygofer opening (fig. 24). Caudodorsal beak in dor-
sal view (fig. 29) truncate at apex. Clasper (fig. 25) al-
most straight and directed posteriad. Apical part of
clasper forming a distinct and angularly protruding
dorsal crest, giving the clasper a humped appearance.
Clasper hollow fairly short. Basal part of clasper grad-
ually widening, not forming an angular clasper heel.
Aedeagus (fig. 27) with fairly long and slender apical
part, but much shorter than in most other species of
this group, and well developed lateral lobes. Aedeagus
from behind (fig. 28) very slender, aedeagal pore nar-
row, oval-shaped and truncate at apex.
Measurements: Body length d: 14.7-17.8 mm (x
15:9) mnt 0) Le SSN Ve van (Ge IUD, ME
0.5); tegmen length d: 17.1-19.1 mm («18.1 mm +
0.8), 2: 21.9-22.3 mm (x 22.1 mm + 0.9); head
length d: 1.2-1.5 mm (x1.3 mm), 2: 1.5-1.9 mm (x
1.7 mm); pronotum length d: 1.9-2.3 mm (x 2.1
mm), 2: 2.6-2.9 mm (x2.8 mm); mesonotum length
3: 2.9-3.5 mm (x 3.2 mm), 9: 4.1-4.6 mm (x 4.4
mm); head width 6: 2.5-3.7 mm (x3.3 mm), 9 : 4.0-
4.3 mm (x4.2 mm); width of pronotal collar à : 4.3-
5.2 mm (x4.6 mm), 2: 5.9-6.4 mm (x6.2 mm).
Distribution (fig. 2). — B. gibberosa is recorded
from Popondetta on the Papuan Peninsula and from
Normanby Island. i
Etymology. — Gibberosus (Latin) humpbacked re-
fers to the hump-shaped dorsal protrusion of the
clasper.
170
Baeturia retracta sp. n.
(figs. 2, 32-39)
Holotype: ‘PAPUA NEW GUINEA / Tabubil, Western
Province / 5°15’ S 140° 13’ E/ 13 oct. 1992 / R.B. Lachlan’
(print, day and month written); ‘On loan from / M.S.
Moulds / Sydney’ [print, violet label], & , AMS). — Paratypes:
same data as holotype but 17.x.1992, 16, Moul; same data
but 29.x1.1992, 16, Moul; Garaina, 11-14.vii.1969, J.L.
Gressitt, 16, BPBM; Gurakor, Wampit R. Valley, 45 mi
from Lae, 670 m, 8.v.1959, L.J. Brass, Sixth Archbold
Exped. to Papua New Guinea, 12, AMNH; Gurakor,
7.vii.1965, H. Pyka, 1d, SMN; Iongai, 1700-1900 m,
9.xi.1965, J. & M. Sedlacek, 19, BPBM; Iongai, 10 km E of
Mt. Albert Edward, 1450-1800 m, 8-10.xi.1965, J.
Sedlacek, 14, BPBM; Kiunga, 18.vii.1970, O.K. McCaw,
1d, AMS; Kiunga, 5° 15’ S 141° 05’ E, 2280 m, 25.v.1970,
O.K. McCaw, 12, Moul; Kiunga, Fly River, 35 m,
viii.1969, J. Sedlacek, 12, BPBM; Koitaki, 1500 ft, x-
xi.1928, Pemberton, 1d, BPBM; Wau, Morobe Distr., 1050-
1100 m, 15.xii.1961, J., J.H. & M. Sedlacek, 16, BPBM;
Yagaum, Madang, 3-5.vi.1965, H. Pyka, 1d, SMN; MANUS:
Manus isl., 24.1.1966, 1d, SMN.
B. retracta can be recognized by its erect and re-
tracted claspers, which lie for the greater part con-
cealed within the pygofer. The species is closely relat-
ed to B. laminifer Blöte, sharing a very similar clasper
with a similarly square, laminiform, dorsal protru-
sion. This protrusion, however, is completely con-
cealed within the pygofer, and therefore hardly visible
in B. retracta. B. retracta is regarded as a separate spe-
cies, since the retraction of the clasper appears to be a
constant character. Furthermore, B. retracta tends to
have a more strongly swollen postclypeus, while males
have a slightly shorter operculum and a shorter, more
rounded, protuberance on the lateral lobe of the py-
gofer.
Description
Body of males light yellow-brown or greyish brown
and distinctly speckled with brown. Females more
red-brown and more densely speckled. Female head
and thorax larger than in male. Tegmina of males 1.1-
1.3 x as long as body length, of females 1.2-1.4 x.
Male abdomen 1.2-1.5 x as long as head and thorax,
of females 1.0-1.3 x.
Head: Ochraceous, densely brown speckled.
Postclypeus weakly protruding, almost oblong-
shaped and bluntly rounded anteriorly. Postclypeus
1.4-2.0 x as wide as long. Postclypeus in lateral view
distinctly swollen with strongly convex anterior mar-
gin.
Thorax: Pronotum ochraceous or greyish brown,
with brown speckling concentrated in, often some-
what reddish, medial band and on medial parts of
pronotal lobes. Mesonotum greenish brown, densely
brown speckled. Two brown spots in front of cruci-
form elevation.
Legs: Ochraceous, often reddish towards tarsi, and
weakly speckled. Fore femur with row of three or four
sharply pointed and erect spines.
Tegmina and wings: Hyaline. Veins ochraceous,
and densely set with setae.
Tymbal organ: Four weakly sclerotized parallel
ridges spanning the tymbal from dorsal to ventral
margin. Three intercalary ridges form a lateral band
across tymbal.
Opercula: Male operculum (fig. 39) very small and
not covering tymbal cavity. Distal part of operculum
erect and reaching to about half-length of meracan-
thus, but not reaching to abdomen. Lateral margin of
distal part short, concavely bent into crest around dis-
tolateral corner of basal part, and angularly bent into
long and concave distal margin. Distomedial corner
rounded, medial margin straight. Male from longai
with larger, more oval-shaped, operculum. Female
operculum (fig. 38) almost as large as that of male,
broadly sickle-shaped and erect, with almost continu-
ously rounded distal margin.
Abdomen: Male abdomen not inflated, ochraceous
brown and brown speckled, though ventrally un-
speckled. Segmental hind margins sometimes red-
dish. Tergites 3-6 almost transparent laterally. A dis-
tinct lateroventral row of brown spots on tergites 3-8.
First tergite very short and medially almost complete-
ly hidden under metanotum. Female abdomen more
robust than that of male, dark reddish brown, and
densely speckled. Lateroventral spots on tergites 3-8
very distinct. Ovipositor sheaths not reaching to apex
of caudodorsal beak. Female caudodorsal beak in dor-
sal view (fig. 32) truncate at apex (pointed in longai
specimen).
Male genitalia: Pygofer in lateral view as in fig. 33.
Dorsal margin concave, but convexly bent into al-
most straight caudodorsal beak and concave near apex
of beak. Distal margin almost straight, but concavely
bent into straight margin of beak. Protuberance on
lateral lobe of pygofer bluntly rounded and weakly
projecting posteriad. Pygofer lobe sometimes forming
a small rounded corner just above this protuberance.
Ventral margin almost straight, but convexly bent
just under lateral protuberance and weakly concave
near base. Ventral margins converge to sharp angle at
base of pygofer opening (fig. 36). Caudodorsal beak
in dorsal view (fig. 37) weakly bicuspidate at apex.
Clasper strongly upcurved, reaching into basal part of
caudodorsal beak. Dorsal part of clasper with lamini-
form and square-shaped protrusion. Clasper hollow
fairly short. Aedeagus (fig. 35) with long apical part,
but shorter and broader than in most other species of
this group, and with well developed lateral lobes.
Basal part of aedeagus very stout. Aedeagus from be-
hind (fig. 34) long and slender, though fairly broad
between its lateral lobes. Aedeagal pore narrow, oval-
De Boer: The Baeturia nasuta group
shaped and truncate at apex.
Measurements: Body length d: 14.9-19.6 mm (x
17.2 mm + 1.4), 2: 16.3-18.1 mm (x 17.5 mm +
0.7); tegmen length d: 18.2-21.9 mm (x 20.6 mm +
IMD) 222174225 95mm &23.0"mm 3a) ahead
length d: 1.3-1.5 mm (x 1.4 mm), 2: 1.6-1.8 mm (x
1.7 mm); pronotum length d: 2.1-2.5 mm (x 2.2
mm), 9: 2.4-2.7 mm (x2.6 mm); mesonotum length
3S: 3.2-4.1 mm (x 3.7 mm), 9: 3.9-4.8 mm (x 4.2
mm); head width d: 3.4-3.8 mm (x3.7 mm), © : 3.9-
4.2 mm (x 4.1 mm); width of pronotal collar 4 : 4.7-
5.6 mm («5.1 mm), 2: 5.7-6.3 mm (x5.9 mm).
Distribution (fig. 2). — B. retracta is widely distrib-
uted in Papua New Guinea, and also recorded from
Manus Island of the Admiralty Islands.
Etymology. — Retracta refers to the retracted clasp-
ers of this species.
ACKNOWLEDGEMENTS
For the loan of material I am indebted to: Dr R.T.
Schuh (AMNH); Mr. B.J. Day (AMS); Dr W.J. Knight
and Mr. M.D. Webb (BMNH); Mr. G.M. Nishida and
Mr. K. Arakaki (BPBM); Mr. J. van Tol (RMNH); Mr.
F. Heller (SMN); and to Mr. M.S. Moulds for the
loan of specimens from his private collection.
I would like to thank Mr. G. Verlaan for technical
assistance and and Mr. D.A. Langerak for preparing
the map (fig. 2). I am indebted to Prof. Dr F.R.
Schram and Dr J.P. Duffels (Institute for Systematics
and Population Biology, University of Amsterdam)
for their critical reading and comments on the manu-
script.
REFERENCES
Anonymous, 1938. Atlas van tropisch Nederland: [i-ix],
maps 1-31b, legenda 1-31b, 1-17 (Koninklijk Neder-
lands Aardrijkskundig Genootschap & Topografische
Dienst in Nederlandsch-Indié, Amsterdam/Batavia).
Anonymous, 1966. List of New Guinea localities: 1-19
(Bernice P. Bishop Museum, Honolulu).
Anonymous, 1968. The Times atlas of the world: i-xliii, 1-
272, pls. 1-123. (Times Newspapers Ltd., London).
Blôte, H. C., 1960. The genus Baeturia Stal as represented
in New Guinea (Homoptera, Cicadidae). — Zoologische
Mededelingen, Leiden 37: 61-80.
Boer, A. J. de, 1982. The taxonomy and biogeography of the
nasuta group of the genus Baeturia Stal, 1866
(Homoptera, Tibicinidae). — Beaufortia 32 (4): 57-78.
Boer, A. J. de, 1986. The taxonomy and biogeography of the
conviva group of the genus Baeturia Stal, 1866
(Homoptera, Tibicinidae). — Beaufortia 36 (7): 167-182.
Boer, A. J. de, 1989. The taxonomy and biogeography of the
bloetei group of the genus Baeturia Stal, 1866
(Homoptera, Tibicinidae). — Beaufortia 39 (1): 1-43.
Boer, A. J. de, 1990. Aedeastria, a new cicada genus from
New Guinea, its phylogeny and biogeography
171
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
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the taxonomy of the New Guinean Tibicinidae. —
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Boer, A. J. de, 1992. The taxonomy and biogeography of the
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(Homoptera, Tibicinidae). — Bijdragen tot de Dierkunde
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Boer, A. J. de, 1994a. The taxonomy and biogeography of
the loriae group of the genus Baeturia Stal, 1866
(Homoptera, Tibicinidae). — Tijdschrift voor Entomolo-
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Daly, M. C., M. A. Cooper, I. Wilson, D. G. Smith, & B. G.
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(Homoptera: Cicadidae). — Pacific Insects Monographs
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Duffels, J. P., 1986. Biogeography of Indopacific
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Received: 13 January 1994
Accepted: 11 April 1994
ROSSER W. GARRISON
Los Angeles County Agricultural Commissioner's Office, U.S.A.
A REVISION OF THE NEW WORLD GENUS
ERPETOGOMPHUS HAGEN IN SELYS
(ODONATA: GOMPHIDAE)
Garrison, R. W., 1994. A revision of the New World genus Erpetogomphus Hagen in Selys
(Odonata: Gomphidae). — Tijdschrift voor Entomologie 137: 173-269, figs. 1-230, tabs. 1-4
[ISSN 0040-7496]. Published 15 December 1994.
This revision of adult Erpetogomphus includes a phylogenetic assessment of all 21 species using
outgroup comparison and parsimony algorithm, descriptive biogeography, keys to both sexes,
synonymies, descriptions, type designations, and illustrations, including distribution maps of all
species. Six new species are described: E. agkistrodon, E. leptophis, E. elaphe, E. liopeltis, E. bothrops,
E. heterodon. Erpetogomphus coluber is considered a junior synonym of E. compositus, E. natrix is
considered a subspecies of E. lampropeltis, and a neotype is designated for E. cophias. Phylogenetic
assessment of 41 mostly somatic characters shows Erpetogomphus to be partitioned into three
monophyletic groups: 1) six dark green species (E. constrictor, E. sabaleticus, E. tristani, E. agkistro-
don, E. schausi, E. ophibolus) with mostly allopatric or parapatric distributions along the eastern
coast of Mexico south into northern Colombia and Venezuela, 2) two species (E. leptophis, E. eu-
tainia) with distributions from southern Texas south through Mexico and up the west coast to
Michoacan states, and 3) 13 remaining species (E. elaphe, E. elaps, E. liopeltis, E. bothrops, E. vi-
perinus, E. designatus, E. sipedon, E. lampropeltis, E. crotalinus, E. heterodon, E. compositus, E. boa,
E. cophias) with distributions in the central United States south through Mexico to Costa Rica.
Derived characters states were gleaned mostly from primary and secondary genitalic characters
(head structure, penis, hamules, caudal appendages, vulvar lamina); but satisfactory resolution of
terminal clades, especially of the third group, is difficult due to apparent character reversals.
Dr. R. W. Garrison, Los Angeles County Agricultural Commissioner’s Office, 3400 La Madera
Ave., El Monte, CA 91732, USA
Key words. — Odonata; Gomphidae; Erpetogomphus, systematics; keys; cladistics; phylogeny;
biogeography.
CONTENTS Erpetogomphus agkistrodon sp. n. ....... 199
Erpetogomphus schausi Calvert … 200
Introductory part Erpetogomphus eutainia Calvert … … 201
Introduction nennen ee 174 Erpetogomphus leptophis sp. n... 204
SON tate UE Mal width 174 Erpetogomphus elaphe sp. n. .... uu 205
The status of the name Erpetogomphus elaps Selys … … … … nnen 207
Erpetogomphus menetriesii (Selys) … … … … … 174 Erpetogomphus liopeltis sp. n. nennen 210
BiOlO SAR AE ee re ste 176 Erpetogomphus bothrops sp. n. nennen DIV
Phylogenetics and biogeography Erpetogomphus viperinus Selys … … 215
Phylogeneticanalysis #1. 17/7 Erpetogomphus designatus Hagen in Selys .... 217
EES Er ane el Lure, 180 Erpetogomphus sipedon Calvert … 221
Distuibutionypatternsman. rn... nenne 181 Erpetogomphus lampropeltis Kennedy .......... 223
Déscripiyebioscosraphye rina 182 EA vlampropelis Wenmed yee nn 225
Chace are nel mn 182 E .l. natrix Williamson and
Erpetogomphus species groups … annen 185 WillAmMSsonista tin pme 226
Systematic part Erpetogomphus crotalinus (Hagen in Selys) .. 227
Introduction ern ee ol 185 Erpetogomphus heterodon sp. n... 230
Genus Erpetogomphus Hagen in Selys, 1858 186 Erpetogomphus compositus Hagen in Selys ... 232
bene stillen 188 ErpetogomphuslboasSelys me 236
Erpetogomphus constrictor Ris … n 193 Erpetogomphus cophias Selys … … n 238
Erpetogomphus sabaleticus Williamson … … 195 Acknomledsemn ents 240
Erpetogomphus tristani Calvert … 196 References sth. sen. sa 241
Erpetogomphus ophibolus Calvert ................. 197 Illustrations | ati. et ee 247
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
INTRODUCTORY PART
Introduction
No other group of Middle American Gomphidae
has been in need of revision as much as the genus
Erpetogomphus Hagen in Selys. The last species were
described by Williamson and Williamson (1930),
and although several odonatists have since received
species which are possibly undescribed, there has been
a hesitancy to describe them because so many species
are rare or poorly known. Several years ago, I collec-
ted a few specimens of one species I thought new, but
I realized that an examination of most of the types
would be necessary before I could resolve the pro-
blem.
This paper is the result of those investigations. My
purpose here is to: 1) describe and figure all species
currently going under the generic name
Erpetogomphus, 2) select lectotypes and establish type
localities, if possible, and 3) construct a cladogram of
all the known species so that phylogenetic relation-
ships and distribution patterns can be adduced for the
genus. I have examined types for all taxa except for
two species, E. cophias (type lost), and E. ophibolus.
The last species is distinct, and no confusion exists
about its identity or type deposition. A neotype is se-
lected for E. cophias. I also provide keys for all species.
Twenty-three names (table 1) have been included
in Erpetogomphus, but three, E. severus Selys,
Herpetogomphus rupinsulensis Walsh, and H. pictus
Needham (= Ophiogomphus rupinsulensis (Walsh)
have been transferred to Ophiogomphus. Another, E.
menetriesii (Selys) I consider to be a nomen dubium,
for reasons detailed below; and six new species are de-
scribed. One species thought lost, £. boa Selys, has
been rediscovered, and £. viperinus auctorum is really
an undescribed species. I suspect that several new spe-
cies will be found in Mexico and Central America,
and I hope this paper will make it easier for others to
describe those new species.
The difficulties I have encountered during this
study have mostly been due to the paucity of speci-
mens of various species which have been considered
rare. I have successfully associated both sexes of all
current species except the male of E. agkistrodon,
which is unknown. My task of recording aspects of
intraspecific variation has been made considerably
easier by various museum authorities and individuals
listed under acknowledgements.
History
The genus Erpetogomphus was described (as a sub-
genus) by Selys (1858) to receive two previously de-
scribed species, Gomphus menetriesii Selys, 1850, and
Ophiogomphus crotalinus Hagen in Selys (the latter
originally described from a pair in 1854), as well as
174
two new species, Erpetogomphus elaps Selys, and E.
cophias Selys. In the ‘Appendice’ of the same paper,
Hagen described £. compositus and E. designatus. Selys
described £. boa in 1859 and E. viperinus in 1868.
Some of these species were briefly redescribed or dis-
cussed by Selys and Hagen in 1859, 1869, 1873, and
1878. In 1879, Selys redefined the subgenus
Herpetogomphus (an unjustified emendation of
Erpetogomphus, see Cowley, 1934), under which were
included eight species: E. compositus, E. designatus, E.
viperinus, E. menetriesit, E. elaps, E. boa, E. cophias,
and £. crotalinus. Calvert (1899) redescribed E. vipe-
rinus (now E. bothrops sp. n.) based on material col-
lected in Tepic, and included a diagnosis of the fema-
les of E. viperinus and E. elaps. Kirby (1890), in a
synonymic catalogue of the world Odonata, designa-
ted Herpetogomphus crotalinus as the type species of
the genus. Calvert (1905) included a synopsis for all
Erpetogomphus from Middle America and described
E. eutainia, E. ophibolus, E. sipedon, and E. diadophis.
Calvert (1912a) added £. tristani. Ris (1917) descri-
bed E. constrictor and redescribed what he thought to
be the second known male of £. boa. In 1918
Williamson described E. sabaleticus from a pair from
Colombia, and Kennedy (1918) described E. lampro-
peltis from California. Calvert (1919) described £.
schausi from a male from Guatemala. Finally,
Williamson and Williamson (1930) described two
new species from Baja California, E. coluber and E.
natrix. In the same paper, they presented a synopsis of
what was known for all species. Their summary mak-
es clear that only a few common species were repre-
sented in collections, and that many others were
poorly known and/or described from insufficient ma-
terial.
The status of the name Erpetogomphus menetriesii
(Selys)
This name was first applied by Selys to an incom-
plete male supposedly from Brazil. Selys (1850) brief-
ly compared it to Paragomphus genei, as follows: ‘Its
colouration yellow, almost without spots, prevents
confusion of this species with any other in Europe,
but it resembles in this respect G. pallidus (Ramb.) of
Southern America whose stature is stronger, and
Gomphus menetriesii (new) from Brazil, but the feet of
the last are mostly black, the d abdomen is bordered
by black on the two sides, etc...’. The same specimen
was redescribed it in 1854 as follows: “Head yellow,
except for space between ocelli, that between the eyes
and ciliated margin of the occiput black. Front of
thorax yellow, with 2 median, contiguous stripes, al-
most obliterated, pale red, antehumeral and humeral
stripe equidistant.
3 Appendages lacking. From: Brazil?’ The type is
said to be in the ‘Musée de St- Pétersburg’.
Table 1. Species-group names used in the genus Erpetogomphus
Name
agkistrodon
boa
bothrops
coluber
compositus
constrictor
cophias
crotalinus
designatus
diadophis
elaphe
elaps
eutainia
heterodon
lampropeltis
leptophis
liopeltis
menetriesii
montanus
natrix
ophibolus
pictus
rupinsulensis
sabaleticus
schausi
Severus
sipedon
tristani
viperinus
Original
Genus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Ophiogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Ophiogomphus
Herpetogomphus
Erpetogomphus
Erpetogomphus
Herpetogomphus
Herpetogomphus
Erpetogomphus
Erpetogomphus
Ophiogomphus
Erpetogomphus
Erpetogomphus
Erpetogomphus
Original
Reference
This paper
Selys 1859
This paper
Williamson &
Williamson 1930
Hagen in
Selys 1858
Ris 1917
Selys 1858
Hagen in
Selys 1858
Hagen in
Selys 1854
Calvert 1905
This paper
Selys 1858
Calvert 1905
This paper
Kennedy 1918
This paper
This paper
Selys 1854
Selys 1878
Williamson &
Williamson 1930
Calvert 1905
Needham 1897
Walsh 1862
Williamson 1918
Calvert 1919
Hagen 1874
Calvert 1905
Calvert 1912
Selys 1868
Type locality
Jalapa, Mexico
Vera Cruz, Mexico
Rio Otapa, Veracruz
State, Mexico
San José de Comandu,
Baja Calif., Mexico
Pecos River, Texas;
corrected to vicinity
of Roswell, NM'
Misantla, Veracruz,
Mexico
Mexico
Mexico
Pecos River, Texas;
corrected to vicinity
of Roswell, NM!
Texas
Costa Rica
Mexico
Guerrero, Mexico
Aragon, New Mexico
Fillmore, California
Blue Creek, Belize
La Estanzuela, Nuevo
Leon, Mexico
Brazil?
Yellow Town, Montana
San Jose de Comandu,
Baja Calif., Mexico
Atoyac, Veracruz,
Mexico
Ithaca, New York
Rock Island, Illinois
Cristalina, Colombia
Purulta, Guatemala
Colorado
Guadalajara, Mexico
Oricuajo, Costa Rica
Orizaba, Veracruz,
Mexico
GARRISON: Revision of Erpetogomphus
Location
of type
UNAM (8)
IRSN (6)
USNM (8)
UMMZ (d)
MCZC (9)
SMF (8)
MNHP (d)
MCZC (8)
MCZC (8)
BMNH (ó)
FSCA (d)
MNHP (d)
BMNH (d)
USNM (8)
USNM (d)
FSCA (8)
FSCA (d)
Lost
IRSNG (8)
UMMZ (d)
BMNH (d)
CUIC (8)
Lost
UMMZ (8)
ANSP (8)
Lostd (d, 2)
BMNH (9)
ANSP (d)
IRSN (8)
Type status
Holotype
Lectotype
Holotype
Holotype
Holotype
Lectotype
Neotype
Lectotype
Lectotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Holotype
Lectotype
Reference for
first placement
in Erpetogomphus
This paper
Selys 1859
This paper
Williamson &
Williamson 1930
Hagen in Selys
1858
Ris 1917
Selys 1858
Hagen in Selys
1858
Hagen in Selys
1858
Calvert 1905
This paper
Selys 1858
Calvert 1905
This paper
Kennedy 1918
This paper
This paper
Selys 1858
Selys 1879
Williamson &
Williamson 1930
Calvert 1905
Needham 1897
Walsh 1862
Williamson 1918
Calvert 1919
Selys 1878°
Calvert 1905
Calvert 1912
Selys 1868
Present
placement
E. agkistrodon
E. boa
E. bothrops
E. compositus
E. compositus
E. constrictor
E. cophias
E. crotalinus
E. designatus
E. eutainia
E. elaphe
E. elaps
E. eutainia
E. heterodon
E. I. lampropeltis
E. leptophis
E. liopeltis
Nomen dubium
Ophiogomphus
Severus montanus
E. lampropeltis
natrix
E. ophibolus
Ophiogomphus
rupinsulensis
Ophiogomphus
rupinsulensis
E. sabaleticus
E. schausi
Ophiogomphus
Severus
E. sipedon
E. tristani
E. viperinus
! See remarks under Z. designatus.; * Transferred to Ophiogomphus by Needham (1899); * Transferred to Ophiogomphus by Hagen (1874)
' As Herpetogomphus; ° Transferred to Ophiogomphus by Selys (1879)
175
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
In 1858, Selys discussed £. menetriesii under E. cro-
talinus (in translation):
‘N.B. In the Synopsis des Gomphines, I described
under the name of Ophiogomphus Menetriesii an in-
complete male individual, which was communicated
to me a long time ago by the knowledgeable M.
Menetries. Now, I am inclined to believe that this is
not a species distinct from crotalinus.
One reason that made me suppose the species dif-
ferent, was its occurrence in Brazil, but it was still
necessary that this fact be duly confirmed and that the
missing anal appendages be known.
This 8 is a little smaller than our specimens [the _
presently described males and female of E. crotalinus],
the space around the ocelli is distinctly black, the ap-
pearance of the reddish antehumeral stripes more
closely approaches the median [stripes], the humeral
is more distinct and more elongated, the base of the
abdomen is less swollen, the dorsal yellow stripe of
3rd, 4th, and 5th segments seems modified: it con-
sists of spots of three lobes [each], whose posterior
isn't visibly narrowed into a little round head. (This
note is based on diagnosis and a description made a
long time ago, so that I have not been able to compare
the specimen since I have had crotalinus at hand.)
A female, same indication of origin [Brazil] from
the Mus. St. Petersburg, communicated to M. Hagen
by M. Menetries differs very slightly from crotalinus
by the following: 6 cells under the pterostigma.; costa
yellower, tarsi black, the first article [tarsomere] of the
posterior [leg] yellow, black tibiae bilineated with yel-
low outside; femora yellow with a double black exter-
nal line shorter and finer at the ends. Abdomen thick-
er, marked with black; 9th segment black above with
a dorsal round spot, touching the posterior margin.
[Abdominal segment] 10 and appendages light red,
the end of these last [structures] pale not marked with
black [italics are Selys]. Vertex more black.
Dimensions a little more robust.
Not being certain of its identity, I fear to cause
confusion later by putting the name E. menetriesii in
true crotalinus from Mexico, even though I have al-
ready published this name [menetriesii] in speaking of
another species (page 102 Revue des Odonates, 1850;
and Synopsis des Gomphines, 1854).
If its identity is confirmed, it would be necessary to
replace the name of E. crotalinus with that of E.
Menetriesii. .
Finally, Selys (1878) described Herpetogomphus
menetriesii based on ‘un mâle in complet [in MNHP]’
and female [in 1RSN] from Guatemala. He states: ‘I
believe it identical with the d example described in
Synopsis No. 20 [1854] and reported with doubt as
crotalinus in the Monographie [1858]. Calvert
(1905) pointed out the inconsistencies of the three
descriptions and left open the question of what E.
176
menetriesii really is.
According to the original description (Selys, 1854),
E. crotalinus cannot be E. menetriesii. A supplemen-
tary male of £. crotalinus described in 1858 (in ZMHB,
examined) is unusual in having faint antehumeral
stripes, but no £. crotalinus I have examined fits the
description of E. menetriesii of 1854 (with middorsal,
antehumeral and humeral stripes).
The 1854 description is too brief to indicate what
E. menetriesii is, and I cannot associate it with any
known species. According to Hagen (1861), it was
destroyed. I have been unable to trace the 1858 fe-
male described by Selys, but I was able to examine the
‘pair described in 1878. The two specimens, which
have determination labels by Selys, are really two fe-
males of E. eutainia Calvert. However, I cannot be
certain that these females are conspecific with Selys’
original description of E. menetriesii. Selys and others
apparently had difficulty in associating specimens of
E. eutainia, because a male from Texas in the IRSN is
given the manuscript name of Erpetogomphus berus,
another female is identified as Cyanogomphus ?mexic-
anus (by R. Martin), and, of course, the two females,
one in the IRSN, the other in the MNHP, are labelled as
E. menetriesit.
Thus, the original description of E. menetriesii may
refer to E. eutainia, but because no one can ever be
sure of this, I propose that Æ menetriesii should be
considered a nomen dubium.
Biology
Little is known of the biology of the genus, though
their habits as recorded in the literature mirror the be-
haviour of other Gomphidae. Adults are most com-
monly found near shores of streams and rivers, but
they may also be found in agricultural stubble or on
tree branches near streams. Many species are seldom
encountered and only a few species appear to be com-
mon in collections.
Available notes taken from the literature, personal
correspondence, and personal observations cited
under various species accounts indicate that female
Erpetogomphus oviposit by swiftly tapping the water’s
surface or by hovering motionless over moderately
swift water systems. Details of copulation and copula-
tion time are largely unknown, although the bizarre
male and especially female morphological adaptations
for achieving the tandem position in E. tristani have
been detailed by Calvert (191 2a).
Larvae of E. designatus, E. compositus, and E. lam-
propeltis were keyed by Needham and Westfall
(1955), and illustrations and full descriptions of £.
crotalinus and E. lampropeltis natrix were given by
Novelo and Gonzalez (1991). The larva of E. sabalet-
icus was recently described and illustrated by Belle
(1992).
GARRISON: Revision of Erpetogomphus
Table 2. Characters used in cladistic analysis of Erpetogomphus adults. & (male) or 2 (female) in parentheses indicates sex to
which character pertains. An asterisk (*) indicates an ordered multistate character. Numbers in parentheses following char-
acter states refer to figure numbers.
Number/character Primitive state Derived state(s)
1. Postocciput width (2) tumid, convex linear
2. Postocciput (9) not visible from above visible from above
3. Occiput (2) semicircular transverse
4. Vertex (2) with long median trough with no trough
5. Postocellar ridge (2) incomplete complete
6. Vertex: sexually dimorphic not sexually dimorphic
7. Frontoclypeal suture with black with no black
8. Hind margin of metepimeron with black with no black
9. Hind wing 3-celled anal loop present absent
10. Anterior hamule* (2) states 1-5 (fig. 208) state 0 (fig. 208)
11. Posterior hamule* (4) states 1-4 (fig. 209) state 0 (fig. 209)
12. Tip of posterior hamule* (4) states 1-5 (fig. 210) state 0 (fig. 210)
13. Posterior lobes of penis segm. 1 (4) small well developed, divided lobes
14. Penis: segment 3 dorsally (3) with pair of tubercles lacking tubercles
15. Penis: segment 4 (4) about twice as long as wide (fig. 211) about as long as wide (fig. 211)
16. Prepuce of penis* (d) states 1-3 (fig. 212) state 0 (fig. 212)
17. Lateral lobe of penis-shape* (4) states 1-5 (fig. 213) state 0 (fig. 213)
18. Membranous hood of penis (à) not overlapping distally overlapping
19. Shape of cornua of penis* (4) states 1-6 (Fig. 214) state 0 (fig. 214)
20. Development of cornua of penis* (à) reduced (1), absent (2) present (fig. 214)
21. Length of vulvar lamellar plates (9) about 0.25 length of sternite > 0.50 length of sternite
22. Medial margins of vulvar lamina (9) folded under not folded under
23. Postlamellar ridge (9) Y-shaped semicircular
24. Postlamellar ridge with juncture (9) posterior to plates (e.g. figs. 184, 195) at or before margin of plates (fig. 192)
25. Ventral base of cercus (6) with large blunt tooth with no tooth (fig. 94).
26. Ventral base of cercus (4) with no carina with a carina
27. Distal part of cercus ventrally (4) with no carina with a carina
28. Dorsal surface of cercus (4) with a tubercle without a tubercle
29. Ventral surface of cercus (d ) linear or concave convex
30. Curvature of epiproct with tips (4) about parallel to base at about 90° to base
31. Tips of epiproct (4) spatulate or bidentate pointed
32. Penis segment 3 (3) short and stocky longer than wide
33. Penis guard (d) quadrate in cross-section circular in cross-section
34. Epiprocts (3) approximate and curved at distal 0.50 not approximatebarely curved
35. Spermatheca (9) states 1-2 (fig. 215) state 0 (fig. 215)
36. Dorsal surface of epiproct (4) with dorsal spine (fig. 95) unarmed
37. Ventral base of cercus (3d) with pebble-like carina otherwise (fig. 97)
38. Membranous hood of penis (4) partially (1) or wholly (2) completely covering subcuticular membrane
exposing subcuticular membrane
39. Ventral base of cercus (4) with carina ending in tooth (fig. 116) with carina (if present) not ending in tooth
40. Postocciput (9) with lateral depressions (fig. 165) with no lateral depressions
41. Occiput (9) with a posteriorly directed medial without a spine
spine fig. 151
PHYLOGENICS AND BIOGEOGRAPHY
Phylogenetic analysis
I have attempted to construct a cladogram based
on an algorithm which infers phylogenies based on
the presence of shared derived characters. Using the
outgroup comparison method of Watrous and
Wheeler (1981), I consider Ophiogomphus to be the
outgroup to Erpetogomphus, following Carle (pers.
comm.) in his assessment of gomphid dragonfly clas-
sification. In this analysis, I have used the most austr-
al species, O. severus Hagen.
The genus Erpetogomphus has never been easily de-
fined (Calvert 1905, Needham and Westfall 1955),
these authors using a series of characters to character-
ize the genus. A synapomorphy apparently unique to
Erpetogomphus is the condition of the epiprocts of the
males: all have gently (at least 110°) to strongly dor-
sally curved, non-divergent epiprocts.
Carle (1986), in his treatment of the higher classifi-
cation of the Gomphidae, included Erpetogomphus in
the tribe Onychogomphini of the subfamily
Onychogomphinae. In his key to the eight subfami-
lies of Gomphidae, he characterized the
Onychogomphinae as follows: 1) hind femur shorter
than head, 2) anal triangle typically 4-celled with
small rectangular cell along inner margin, 3) anterior
hamuli with shoulder and end hook forming lateral
u-shaped notch, and, 4) female sternum 9 with large
U- or V-shaped basal membranous area. Within this
subfamily, the tribe Onychogomphini (including
Erpetogomphus) is distinguished from the tribe
Crenigomphini by the following characters (antago-
nistic characters for Crenigomphini in parentheses):
postgenal suture present (absent), male tibial laminae
WF,
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
present (absent), penile segment one (= peduncle)
without cuplike thin-walled hood (with bilobed cup-
like thin-walled hood), female sternum 9 with dome-
shaped membranous area not extending to distal half
of sternum (this structure extending to distal half of
sternum). None of these characters is stated by Carle
(1986) to be uniquely derived for any of the above
categories. Carle (pers. comm.) and I agree that
Ophiogomphus is the most likely outgroup to
Erpetogomphus, and we agree that both should be
placed in the Onychogomphini. Three characters
seem unique to the New World Onychogomphini,
which include only Ophiogomphus and
Erpetogomphus. 1) rami of male epiprocts contiguous
along all or most of their length, 2) female sternum 8
with a well-defined costate postlamellar ridge separat-
ing the soft membranous area from the posteriorly
sclerotized area (this ridge reduced in Ophiogomphus),
3) presence of a lateral lobe on the ventrolateral mar-
gin of the fourth penile segment (fig. 213) (reduced
in Ophiogomphus, and not specialized in the £. ophib-
olus group). This last character seems to be a unique
synapomorphy for Ophiogomphus and Erpetogomphus.
Carle (1992) further separated Erpetogomphus into
three subgenera, two of which were new:
Calogomphus (type species E. eutainia) and
Erpetocyclops (type species E. ophibolus). Both Carle
(pers. comm.) and I consider Erpetogomphus to be
composed of three monophyletic groups as discussed
below. While several earlier versions of my cladogram
using series of characters and different states generally
support a conservative grouping of the genera into
three monophyletic groups, I do not think it neces-
sary to provide subgeneric rank to those groups.
A suite of 41 characters, 7 of which are multistate
(table 2), was analyzed using the HENNIG86 comput-
er program (Farris, 1988), with the following op-
tions: The character set was first run using the ie*
(guaranteed to find the most parsimonious trees) op-
tion where all characters were ordered and each was
equally weighted (default options). All autapomor-
phies were included from this analysis. The final set
of 41 characters was finally chosen from larger sets,
each with different coding sequences, because this set
seemed to be the most heuristic. The resultant analy-
sis yielded six equally parsimonious trees. A Nelsen
(consensus) tree (length 106, consistency index 0.62)
was then generated from the six trees. The resultant
characters, tree, and matrix are shown in figs. 208-
216, and tables 2-3. As this manuscript was in review,
Jan van Tol kindly ran the same data set on PAUP
3.1.1 for the Macintosh (Swofford 1993) using unor-
dered as well as ordered sequence of multistate char-
acters. His results, which he kindly communicated to
me, resulted in 61 semistrict consensus trees each
with a length of 94 for unordered multicharacter
178
states. His resulting cladogram is similar to that in fig.
216 except that the entire E. crotalinus group (except
for three sister groups: E. elaphe and E. elaps E. sipe-
don and E. lampropeltis and E. boa and E. cophias)
collapses to a polytomious assemblage. (I recorded a
similar cladogram when multistate characters were
listed as unordered for HENNIG86.) When all multis-
tate characters were run as ordered on paup 3.1.1, the
resulting cladogram was the same as shown for fig.
216.
I have not been totally successful in resolving sever-
"al of the terminal clades, due to high degree of appar-
ent homoplasies. The best characters are those of the
secondary genitalia of the male (hamules, penis); but,
despite a thorough examination of these structures, I
am still uncertain of the correct interpretation of
character polarity for several of these complex, mult-
istate character sets (figs. 208-210, 213-214). Little
difficulty was encountered in assigning states to
members of the E. ophibolus and E. eutainia groups,
and their generalized tracks also independently sup-
plement in understanding their phylogeny; but the
same characters in males of the E. crotalinus group
show, if my cladogram is correct, a bewildering com-
plex of character reversals for some characters. The
generalized tracks within the E. crotalinus group (fig.
226) also provide few clues to possible speciation
events, except for that between the £. elapsand E. cro-
talinus (S.s.) groups.
Individual clades are numbered as shown in fig.
216, each of which is discussed below.
Clade 1.1. — E. ophibolus group (six species: E.
constrictor, E. sabaleticus, E. tristani, E. ophibolus,
E. agkistrodon, E. schaust) = subgenus Erpetocyclops
Carle
Character 3. — female occiput semicircular (parallel
development in Clade 1.22112, E. liopeltis)
Character 5. — female postocellar ridge incomplete
(parallel development in Clade 1.211, E. leptophis)
Character 6. — vertex sexually dimorphic (state not
known for male E. agkistrodon, but probably follows
for other members of this clade)
Character 11, state 1. — male with posterior hamule
triangular (fig. 209)
Character 12, state 1. — male with tip of posterior
hamule acuminate (fig. 210)
Character 16, state 1. — male with prepuce well-de-
veloped (fig. 212)
Character 17, state 1. — male with lateral lobe ves-
tigial, broadly angulate with no serrations (fig. 213)
Clades 1.11 (E. constrictor, E. sabaleticus, E.
tristani, E. ophibolus), 1.12 (E. agkistrodon), 1.13
(E. schausi). Clade 1.11 is defined by the following
synapomorphies:
Character 20, state 2 - male with loss of cornuae of
penis (fig. 214). Most Gomphidae (including the
outgroup, Ophiogomphus severus), have a pair of long,
strap-like cornuae. I have hypothesized the character
transformation for this structure throughout the ge-
nus (fig. 214).
Character 25. — ventral base of cercus of male with
large blunt tooth (fig. 94)
Character 28. — dorsal surface of cercus of male
with a tubercle (fig. 94)
I have found no synapomorphy for uniting 1.12
(E. agkistrodon) and 1.13 (E. schausi). The former is
known only from the holotype female, the latter by
two males and a teneral female.
Clades 1.111 (E. constrictor), 1.112 (E. sabaleticus),
1.113 (E. tristanı)
Character 29. — curvature of epiproct of male with
tips completely recurved so as to lie parallel to base
Clade 1.21. — E. eutainia group (two species: E.
eutainia, E. leptophis) = subgenus Calogomphus
Carle
Character 10, state 1. — anterior hamule with
equally divided arms (fig. 52-53, 208) at distal 0.25
Character 11, state 2. — posterior hamule digit-
shaped (fig. 209)
Character 14. — dorsal surface of third segment of
penis of male with two tubercles
Character 16, state 2. — prepuce of male reduced,
largely hidden by frill-like lateral lobe (fig. 212)
Character 17, state 2. — lateral lobe of penis of male
frill-like (fig. 213)
Character 35, state 1. — spermatheca of female a
single tube (fig. 215). I was only able to study this
structure in E. eutainia, as only the allotype of E. lep-
tophis is known. Due to other synapomorphies listed
above, I hypothesize that £. leptophis will possess sin-
gle, rather than paired, spermatheca.
Clade 1.22. — E. crotalinus group (13 species: E.
elaphe, E. elaps, E. liopeltis, E. bothrops, E.
viperinus, E. designatus, E. sipedon, E. lampropeltis,
E. crotalinus, E. heterodon, E. compositus, E. boa, E.
cophias) = subgenus Erpetogomphus Hagen in Selys
Character 16, state 3. — prepuce of penis of male
lacking
Character 35, state 2. — spermatheca of female pai-
red, each forming a long, narrow sac (fig. 215)
Clade 1.221. — E. elaps subgroup (five species: E.
elaphe, E. elaps, E. liopeltis, E. bothrops, E.
viperinus)
Character 17, state 3. — unique shape of lateral lobe
(fig. 213)
GARRISON: Revision of Erpetogomphus
Clade 1.2211 (E. elaphe, E. elaps, E. liopeltis, E.
bothrops)
No satisfactory synapomorphy has been found for
this clade. One character, the spatulate tip of the epi-
proct (fig. 129) separates E. viperinus from its sister
species.
Clade 1.22111 (E. elaphe, E. elaps, E. liopeltis)
No synapomorphy has been found, but the trans-
verse condition of the female postocciput in £. both-
rops and in E. viperinus (fig. 166) split E. bothrops
from the other three sister species.
Clade 1.221111 (E. elaphe, E. elaps)
Character 22. — vulvar lamina of female with mar-
gin folded under ventrally (figs. 180- 181)
Clade 1.222 (E. designatus, E. sipedon, E.
lampropeltis, E. crotalinus, E. heterodon, E.
compositus, E. boa, E. cophias)
No satisfactory synapomorphy has been found to
unite the £. crotalinus group (sensu stricto [s.s.]). The
one aberrant species in the group is £. designatus. It
alone has the following characters which separate it
from the other 7 species: 1) tip of posterior hamule
lacking a tooth (recurrence of this state from the £.
eutainia group) and 2) unique shape and position of
the lateral lobe (fig. 82). The shape of the anterior ha-
mule of E. designatus (fig. 60) is, however, like that of
E. sipedon, E. lampropeltis, E. crotalinus, and E. com-
positus, which thus unites these species into a clade.
The widely distributed £. designatus has been diffi-
cult to place cladistically in this study. It combines
characters of the £. elaps group (for example, linear
condition of lateral lobe and acuminate condition of
the cornua) and even apomorphic characters com-
mon to the £. crotalinus (s.s.) group as noted above.
Clade 1.2222 (E. sipedon, E. lampropeltis, E.
crotalinus, E. heterodon, E. compositus, E. boa, E.
cophias)
Character 12, state 5. — tip of posterior hamule of
male with anteapical tooth
Character 17, state 5. — unique shape of lateral lobe
of fourth segment of penis of male (fig. 213). The
small, semicircular condition of the lateral lobe
present in Clade 1.2222 is similarly developed in
Clade 1.12 (E. schausi).
Clades 1.22221 (E. sipedon), 1.22222 (E.
lampropeltis), and 1.22223 (E. crotalinus, E.
heterodon, E. compositus, E. boa, E. cophias)
No satisfactory apomorphy has been found to di-
chotomize these clades. The distinctive shape of the
cornuae of £. sipedon and E. lampropeltis (figs. 83-85)
has separated these two species from clade 1.22223.
170)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Table 3. Character matrix used in producing Nelsen (consensus) tree shown in Fig. 216.
Character No. 1 D
N
4 5 6
(SS)
Ancestor
Oph. severus
constrictor
sabaleticus
tristani
ophibolus
agkistrodon'
schausi
eutainia
leptophis
elaphe
elaps
liopeltis
bothrops
viperinus
designatus
sipedon
lampropeltis
crotalinus
heterodon
compositus
boa
cophias
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"Only the holotype 2 is known for this species. Male characters were scored as ? (unknown), on 0? or 1?, based on af-
finity with rest of the E. ophibolus group. * Character prevalent in this species. * Basal carina weak in E. boa.
* This structure not examined for this species. Plesiomorphy/apomorphy hypothesized by examination of spermathe-
ca in closely related species, and placement of these taxa in cladogram due to other synapomorphies.
Clades 1.222231 (E. crotalinus) and 1.222232 (E.
heterodon, E. compositus, E. boa, E. cophias)
No satisfactory apomorphy has been found for cla-
de 1.222232. Erpetogomphus crotalinus was separated
because of the unique shape of its cornuae (fig. 86).
Clades 1.2222321 (E. heterodon), 1.2222322 (E.
compositus), 1.2222323 (E. boa, E. cophias)
The unique shape of the cornua (figs. 87-89) has
segregated E. heterodon and E. compositus from clade
1.2222323. Apomorphies for the last clade, repre-
senting the two closely related species, £. boa and E.
cophias, are:
Character 10, state 5. — unique shape of the anteri-
or hamule of male (fig. 208)
Character 11, state 4. — unique shape of the poste-
rior hamule of male (fig. 209)
Character 19, state 6. — unique shape of the cornua
of male (fig. 214)
The clades mentioned above have problems pri-
marily within the E. crotalinus group (sensu lato,
[s.L]), and most dichotomies are based on one or two
highly complex character states of the penis. I realize
that my interpretation of the morphoclines of this
character is open to reevaluation, which could result
in a somewhat different shuffling of some species
180
within the larger group. Only two species in this
group, £. boa and E. cophias, are easily characterized
as a monophyletic group.
A further analysis to resolve some of the difficulties
described above will have to await further specimens
and a proper analysis of their larvae. As stated earlier,
I have not assigned formal names to subordinate
clades, because further investigation may involve
transfer of some monophyletic or paraphyletic groups
to other clades, nor can I see any purpose in a prolife-
ration of infrageneric names.
Cladospecies
Of the 21 species described here, seven (33%) can
be defined by autapomorphies and can be termed cla-
dospecies (Ackery and Vane-Wright, 1984). These
cladospecies with their uniquely derived character sta-
tes are shown below.
Clade 1.112. E. ophibolus. Character 36, dorsal sur-
face of epiproct of male with a dorsal spine (fig. 95)
Clade 1.211. E. leptophis. Character 41, female oc-
ciput with a posteriorly directed medial spine (fig.
151)
Clade 1.212. E. eutainia. Character 37, ventral
base of cercus of male with a pebble-like carina.
Clade 1.2211112. E. elaphe. Character 10, state 3,
m
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unique shape of anterior hamule of male (fig. 208)
Clade 1.22221. E. sipedon. Character 38, state 2,
membranous hood of penis of male wholly exposing
subcuticular membrane.
Clade 1.2222322. E. compositus. Character 38,
state 1, membranous hood of penis of male partially
exposing subcuticular membrane.
Clade 1.22223232. E. cophias. Character 39, ven-
tral base of cercus in male with carina ending in tooth
(fig. 116), and Character 40, postocciput of female
with lateral depression (fig. 165). These two charac-
ters are probably correlated: I hypothesize that the
postoccipital depressions of the female receive the in-
ferior carinal tooth of the cerci of the male during
copulation.
I have been unsuccessful in finding autapomor-
phies for the 14 other species, and they can be termed
paraspecies (Ackery and Vane-Wright, 1984). A fur-
ther, more detailed analysis of these species may yield
autapomorphies. The cladist may question the status
of species recognized by single or combinations of
plesiomorphic characters, but I agree with Ackery and
Vane-Wright (1984: 10), who cogently argue: ‘the
traditional groupings often (but by no means always)
turn out to be good. The cladist must, therefore, al-
ways retain respect for the ‘Gestalt approach (espe-
GARRISON: Revision of Erpetogomphus
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cially when practised by field workers), unless armed
with concrete evidence for some contrary arrange-
ment...
Distribution patterns
Table 4 details the distributional patterns of all
New World Gomphidae according to five political re-
gions: Canada/Alaska, U. S. A., Mexico, Central
America, and South America. All speciose (z.e., N > 3
species) gomphid genera in the New World have pro-
portionally more species in the United States and in
South America than Erpetogomphus. Erpetogomphus
has the most species in Mexico (76%). Central
America follows with 48%, the United States with
24%. No other genus is proportionally as speciose in
Mexico. The Gomphus complex of genera
(Arigomphus, Gomphus and subgenera, Stylurus), and
Ophiogomphus are dominantly North American, and
several genera (Agriogomphus, Aphylla,
Archaeogomphus, Cyanogomphus, Gomphoides,
Neogomphus, Phyllocycla, Phyllogomphoides,
Progomphus, and Zonophora) are South American.
Epigomphus is most speciose in Central America,
where 46% of all described species occur, closely fol-
lowed by South America.
Figure 217 shows the density and approximate
overall distribution of species of Erpetogomphus as
181
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
J
shown by 150 km squares. Of the five species found
in the United States, most are found in the south-
western states. The most northerly distribution for
the genus is Grant County, Washington (46°50’N),
for E. compositus (Paulson and Garrison, 1977). The
greatest Concentration in species is in south-central
Mexico between approximately 18°-20°N with eight
species (31%) of all the total number of species in the
genus found in southern Veracruz alone.
Descriptive biogeography
In this section I describe the current distribution
range (= tracks) of various monophyletic groups ba-
sed on my cladogram. Unfortunately, the numerous
problems associated with the cladistic analysis conco-
mitant with the poorly known or spotty distributions
(most of which may be due to inadequate collecting
in parts of Mexico) preclude a discussion of vicarian-
ce biogeography. However, I do discuss possible spe-
ciation sequences within the E. ophibolus and E. eu-
tainia groups.
The cladogram (fig. 216) for Erpetogomphus di-
vides the 21 known species into three monophyletic
groups. The generalized track for the genus is shown
in fig. 218. The first two groups, the E. ophibolus and
E. eutainia groups, have generalized tracks encom-
passing south Texas, eastern Mexico south into
northern Colombia and Venezuela (figs. 219-220).
The E. ophibolus group comprises six closely related
species united by seven derived character states.
Figure 221 shows that the distribution of all six spe-
cies except E. schausi are largely allo- or parapatric.
Little can be said of E. agkistrodon or E. schausi due to
paucity of records. Erpetogomphus ophibolus occupies
an area between two disjunct populations of £. con-
strictor. Further collecting may show E. constrictor to
be sympatric or parapatric with E. ophibolus. Aside
from E. agkistrodon and E. schausi, the track of E.
ophibolus was probably the first to separate from the
E. ophibolus group, a hypothesis consistent with the
results of my cladogram (fig. 216). The three remain-
ing species, E. constrictor, E. tristani, and E. sabaleti-
cus, are Closely related sister taxa whose tracks are
largely parapatric. Presumably their differentiation
(by allopatric speciation?) took place from a common
ancestor which occupied part of the present range of
all three species.
The tracks of the second monophyletic group com-
prises just two species, the relatively widespread E.
eutainia and the peripheral E. leptophis (fig. 222). The
latter, known only from the holotype and allotype
from Belize, is sympatric with E. ophibolus, but its ge-
nealogical affinities are unquestionably with E. eutai-
nia (fig. 216). Erpetogomphus leptophis probably arose
as a peripheral isolate from E. eutainia, or from a
common ancestor, since it is a more apomorphic spe-
182
cies than £. eutainia.
The remaining 13 species comprise two monophy-
letic subgroups, one predominantly austral, the other
more boreal. Their generalized tracks are shown in
figs. 223-226. The E. elaps subgroup contains five
species, whose somewhat limited distributions occu-
py the western and eastern lowlands of Mexico south
along the mountainous areas into Costa Rica.
Erpetogomphus elaps and E. elaphe are closely related
sister species which are parapatric in Guatemala. A
slight but consistent southern variant of E. elaps occu-
pies the area between more northerly typical popula-
tions of E. elapsand E. elaphe (see discussion under £.
elaps): it may be an incipient species. These two (or
three?) species, like those for E. constrictor, E. tristani,
and E. sabaleticus, probably arose from an E. elaps-like
ancestor or from one another. Interpretations of the
tracks of the remaining three species is problematical
due to the uncertain position of £. /iopeltis in my cla-
dogram. It possesses characters common to £. both-
rops and E. viperinus. No convincing apomorphy
unites £. /iopeltis with the sister taxa £. elaps and E.
elaphe, so its genealogical relationship must remain in
doubt. The two species with the greatest ranges (£.
elapsand E. bothrops) are broadly sympatric; E. liopel-
tis and E. viperinus are allopatric.
The E. crotalinus (s.s.) subgroup comprises the bo-
real group of eight species, with at least one represen-
tative occurring throughout most of the southern half
of the United States. Their complex, mostly overlap-
ping distributions (fig. 226) and questionable phylo-
genetic sequence in the cladogram preclude any
meaningful discussion on their biogeography.
Erpetogomphus boa and E. cophias stand out because
of three synapomorphies. Their limited distribution
and close proximity indicate speciation, possibly by
allopatric means. Erpetogomphus sipedon and E. heter-
odon make up the only other allopatric species pair in
this complex subgroup. Although distribution of £.
heterodon and E. compositus overlap, I have seen no
evidence of these two species collected together.
It is obvious that a more thorough knowledge of
the phylogeny of the genus will be necessary before
interpretation of biogeographical events is possible.
Of particular interest will be a methodological inter-
pretation of the biogeography of the genus, for then
we may be able to determine which species tracks are
the result of dispersal and gene flow or vicariant
events.
Characters
The best diagnostic characters for males are found
in the caudal appendages, accessory genitalia, and
overall body maculation. Most species are easily iden-
tified by the caudal appendages. The cerci (superior
appendages) assume a wide variety of shapes (figs. 92-
GARRISON: Revision of Erpetogomphus
Table 4. Distribution of species of New World Gomphidae. Number under each region = area of region in square miles; %
directly under area = % of total land area; N = total number of species; number under each region = total number of species;
number in parentheses () = percent of total number of species in genus. Phylogenetic classification is after Carle (1986).
Genera are arranged alphabetically under each tribe.
Canada/Alaska U.S.A. Mexico Cent. Amer.” South Amer.
4,438,221 mi’ 3,022,261 mi 761,530 mi 208,800 mi’ 6,597,386 mi
Genus N 29.5% 20.2% 5.2% 1.1% 43.9%
Hageniinae
Hagenini
Hagenius 1 1 (100) 1 (100)
Octogomphinae
Hemigomphini
Neogomphus 3 3(100)
Octogomphini
Lanthus 2 1 (50) 2 (100)
Octogomphus 1 1 (100) 1 (100) 1 (100)
Stylogomphus i 1 (100) 1 (100)
Gomphinae
Gomphini
Arigomphus 7 3 (43) 7 (100)
Dromogomphus 3 1 (33) 3 (100)
Gomphus 38 13 (34) 38 (100) 1 (3)
Stylurus 13 12 (92) 4 (31)
Epigomphinae
Epigomphini
Epigomphus 26 6 (23) 12 (46) 10 (38)
Austrogomphinae
Archaeogomphini
Archaeogomphus 5 1 (20) 1 (20) 5 (100)
Cyanogomphini
Agriogomphus 4 1 (25) 1(25) 3 (75)
Cyanogomphus 6 6(100)
Tibiagomphus 2 2(100)
Onychogomphinae
Onychogomphini
Erpetogomphus 21 5 (24) 16 (76) 10 (48) 1 (5)
Ophiogomphus 18 8 (44) 18 (100)
Lindeniinae
Gomphoidini
Aphylla 19° 3 (16) 2 (11) 3 (16) 18 (95)
Gomphoides 3 3 (100)
Idiogomphoides 2 2 (100)
Peruviogomphus 2 2 (100)
Phyllocycla 31 3 (10) 4 (13) 27 (87)
Phyllogomphoides 43 2 (5) 12 (28) 7 (17) 27 (63)
Lindeniini
Cacoides 1 1 (100)
Melanocacus 2 2 (100)
Mitragomphus 1 1 (100)
Progomphini
Progomphus 59° 4 (7) 9 (15) 7 (12) 52 (77)
Zonophorini
Desmogomphus 2 1 (50) 2 (100)
Diaphlebia 2 2 (100)
Perigomphus 1 1 (100) 1 (100)
Zonophora 10 10 (100)
TOTAL 329 35 (10.6) 97 (29.4) 50 (15.2) 47 (14.3) 180 (54.7)
' Excluding Hawaii; * Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama; * Includes subgenera
Gomphurus, Gomphus, Phanogomphus, Stenogomphurus; * One species, A. caraiba, is endemic to Hispaniola and Cuba, not in-
cluded in N; ” Three species, P. integer, P. serenus, P. zephyrus, are endemic to the Greater Antilles, not included in N.
183
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
116), but they are never longer than segments 9 and
10 combined. All males of the E. ophibolus group, ex-
cept for E. schausi (and possibly the unknown male of
E. agkistrodon) have a dorsal appendage on the distal
third of the cercus. The dorsal surface of the cercus
may be straight as in £. e/aps, strongly curved as in E.
schausi and E. viperinus, angulate as in most species in
the E. crotalinus group, or concave as in the E. boa
and £. cophias group. The inferior third of the cercus
forms a strong carina in E. cophias (weakly so in E.
boa), in the E. crotalinus group, and in E. eutainia. In
the lighter coloured species of the £. crotalinus group,
the carina does not contrast with the rest of the ap-
pendage. A unique series of pebble-like structures
characterizes the basal carina in Æ eutainia. The infe-
rior carina was used by Calvert (1905), although he
wrongly placed E. eutainia among individuals lacking
such a carina. Instead of a basal carina, the £. ophibo-
lus group, E. bothrops and E. viperinus have an inferi-
or carina on the distal third of the appendage. This
structure is weak in E. elaps.
The epiprocts (inferior appendages) are not as var-
ied as the cerci. In most of the E. ophibolus group,
they are strongly curved so that their distal third is
nearly parallel to the basal 0.30. The epiproct of E.
ophibolus is unique in having an anteriorly directed
projection on its dorsal surface near the middle. The
epiprocts are remarkably similar throughout the re-
maining groups: they form a gentle curve with the
tips at approximately 90° to the basal third of the ap-
pendage.
The penes of most species possess a semi-hyaline
pair of flagella or cornuae best seen in dorsal view.
The cornuae are considerably reduced in the £.
ophibolus group (except for E. schausi). Their shape
provides a useful character to differentiate among £.
designatus, E. compositus, and E. lampropeltis. The tip
of each cornua is bluntly rounded in the E. crotalinus
group (except for E. crotalinus). In all others, the tip
forms a sharp point.
The spiny lateral lobes of the penis are well devel-
oped and frill-like in E. eutainia and E. leptophis, pri-
marily knife-like or pointed in E. viperinus, E. hopel-
tis, E. bothrops, E. elaps, and E. elaphe, and a
protruding semicircle in most members of the E. cro-
talinus and E. cophias groups. When the penis is
viewed laterally, the lateral lobe is on edge, because
this structure rotates posteriorly with its flat portion
placed mediodistally. The lateral lobes are poorly de-
veloped in E. constrictor, E. tristani, and E. sabaleticus
(possibly also in E. agkistrodon), but those species and
E. schausi have a unique ventral, arcuate prepuce.
The anterior hamules are always divided, but the
posterior arm in the E. tristani group forms more of a
shoulder than a branch. The bifurcation occupies the
distal 0.25 of the hamule in the E. ophibolus, E. elaps,
184
and E. eutainia groups, the distal 0.50 in the E. cro-
talinus group, and the entire length of the appendage
in E. boa and E. cophias.
The posterior hamules are always more prominent
than the anterior pair. They are broadly triangular in
the E. ophibolus group, but they possess an anterior
shoulder in the £. crotalinus group. In E. eutainia and
E. leptophis, the posterior hamule forms a small fin-
ger-like appendage. All known species except E. desig-
natus, E. elaphe, E. leptophis, and E. eutainia possess a
black apical tooth. This distal tooth is rotated lateral-
ly in E. viperinus and in some E. bothrops.
The best structural characters for separating the fe-
males are those associated with the vertex, occiput,
postocciput (note: my use of the term, ‘postocciput is
not equivalent to the structure immediately surround-
ing the occipital foramen, but, instead refers to the
medioposterior side of the head behind and below the
occipital crest), and vulvar lamina. Calvert (1905)
relied on the curvature of the posterior border of the
occiput, but it is variable within species. I do not use
it here. Better characters are the shapes of structures
on the vertex. In E. constrictor, E. tristani, and E. sab-
aleticus, the median ocellus is placed within a deep
longitudinal trough which accommodates the strong-
ly curved epiproct of the male. In all other species
may be found a transverse trough (figs. 145-146) ora
pair of anterolateral pits (fig. 137) anterior to the me-
dian ocellus. These pits correlate with the shape of the
tip of the male epiproct, and these depressions are al-
so present on the vertex of males. Their shape is use-
ful in distinguishing between the closely related pairs,
E. viperinus and E. bothrops, and between E. composit-
usand E. lampropeltis. A complete postocellar ridge is
present only in all members of the £. tristani group
and E. leptophis. In all others, the middle part of the
ridge is lacking, leaving only postocellar tubercles.
Females of a few species have autapomorphic charac-
ters, such as the raised median area of the occiput of
E. designatus (figs. 156-157), the posteriorly directed
pointed protuberance on the occiput of E. leptophis
(fig. 151) or the postoccipital depressions of E. cophi-
as (fig. 165).
The vulvar laminae vary in the shape of the plates.
The simple horizontal plates are bisected by a small
V-shaped cleft in the £. ophibolus group. The plates
are corrugated and more membranous in £. boa and
E. cophias, often succumbing to much post mortem
distortion. In the £. crotalinus group, each plate is a
large, rectangular, planar structure which meets the
other only at the anterior angle. The resulting median
cleft is only a third or less the width of each plate; but
the cleft is wider and U-shaped in £. designatus. A fur-
ther widening of the cleft is evident in the E. elaps
group. The plates form two small, widely separated,
digit-like structures in E. eutainia and E. leptophis.
A Y-shaped postlamellar ridge is found posterior to
the vulvar lamina. The posterior stem of the Y is ab-
sent in £. eutainia and E. leptophis, so that the ridge
forms a semicircle (figs. 178-179).
I have found no structural differences between the
females of E. heterodon and E. sipedon. Body pattern
must be used to differentiate females of those species.
Erpetogomphus tristani and E. sabaleticus, and E.
elaphe and E. elaps are also closely related pairs, and
their females are distinguishable only by locality.
In well-preserved specimens, body maculation pro-
vides useful characters. Dark thoracic maculation pre-
dominates in the £. tristani group, in E. leptophis, and
in southernmost specimens of E. eutainia. All of those
have a well-defined dark stripe bordering the posteri-
or margin of the metepimeron. Erpetogomphus cophi-
as, E. crotalinus, and most E. elaps have nearly immac-
ulate body patterns. Thoracic maculation is usually
stable, but there is variability in E. lampropeltis, E.
compositus, E. designatus, E. elaps, and E. eutainia.
Two formerly named species, £. natrix and E. coluber,
I consider subspecies of E. lampropeltisand E. compos-
itus, respectively: I have found no morphological
characters which separate those forms. Thoracic pat-
terns in £. elaps are often nearly absent (fig. 13), and
more extensive dark markings characterize popula-
tions of E. eutainia in the north, less extensive in the
south (figs. 7-9).
Abdominal patterns are characterized by a combi-
nation of light and dark banding patterns separated
dorsally by pale longitudinal stripes. The least
amount of black occurs in £. cophiasand E. crotalinus,
the most in £. agkistrodon.
A sharp, prominent, middorsal posterior spine on
abdominal segments 8 and 9 is found in many males
of E. ophibolus, E. bothrops, E. liopeltis, and E. viperi-
nus. The spine is variable and cannot be used reliably
to distinguish among species. For example, many £.
bothrops have no indication of a middorsal spine, but
it is well developed in others. The spines are also
found on abdominal segment 9 in some females of
the E. ophibolus group.
Erpetogomphus species groups
I divide the 21 species into five groups. I believe the
unknown male of one species (E. agkistrodon) will fol-
low those characters listed for that sex discussed be-
low.
1. E. ophibolus group (= subgenus Erpetocyclops
Carle, 1992). Males (probably so for £. agkistrodon,
when discovered): With a distinct curved prepuce,
posterior hamule forming a triangle culminating in a
sharp black tooth. Females: Vulvar lamina simple,
small, relatively undifferentiated postocellar ridge
complete, though curved in E. tristani, E. sabaleticus,
and E. constrictor. Six species: E. tristani, E. sabaleti-
GARRISON: Revision of Erpetogomphus
cus, E. constrictor, E. ophibolus, E. agkistrodon, E.
schaust.
2. E. eutainia group (= subgenus Calogomphus
Carle, 1992). Males: Prepuce small, obscured by lat-
eral lobes of penis; lateral lobes of penis strongly de-
veloped, forming a spiny frill; posterior hamule fin-
ger-like, lacking an apical tooth. Reduced prepuce
present. Females: Vulvar lamina small, finger-like,
widely separated; post-laminar ridge semicircular, not
Y- shaped. Two species: E. leptophis, E. eutainia.
3. E. elaps group. Males: Lateral lobe of penis knife-
like. Females: Vulvar lamina connected anteriorly,
but separated by a cleft 1.0 or more the width of each
plate. Five species: E. bothrops, E. liopeltis, E. viperi-
nus, E. elaphe, E. elaps.
4. E. crotalinus group. Males: Superior surface of
cercus angulate (except for E. compositus); anterior ha-
mule divided at distal half. Females: Vulvar lamina
large, planar, rectangular, meeting at anterior margins
only, resultant cleft one third the width of each plate
(except for E. designatus). Six species: E. crotalinus, E.
lampropeltis, E. compositus, E. heterodon, E. sipedon, E.
designatus.
5. E. cophias group. Males: Anterior hamule talon-
like, divided along its full length; posterior hamule
broadly spatulate, surmounted by a small, black,
curved anteapical tooth. Females: Vulvar lamina di-
agonally corrugated; distal and lateral margins with a
strongly raised semicircular ridge. Two species: E.
boa, E. cophias.
SYSTEMATIC PART
Introduction
Under each species, I have endeavoured to include
a complete synonymy (some minor references may be
lacking for E. designatus, a common species in the eas-
tern United States), a full description, type data, diag-
nosis, remarks, biology, distribution, and material
examined. For new species, I have designated alloty-
pes when the opposite sex was available, but I have
not established allotypes for syntypic material. Under
comments, I have quoted older authors (Ris, Selys,
Hagen), when this information contributes to the his-
tory of the species. For £. agkistrodon, I include a dis-
cussion of what the male should be like, based on my
cladogram. Unless otherwise stated, measurements
are taken from 20 males and 20 females, when availa-
ble, and from as many localities as possible.
Abdominal measurements include caudal appenda-
ges. Full locality data are given for all species, except
for the following species where abundant material
was available: £. eutainia, E. elaps, E. viperinus, E. de-
signatus, E. I. lampropeltis, E. I. natrix, E. crotalinus,
and £. compositus. For these species, I have listed sta-
tes, counties, brief data, and repository. Full locality
185
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
data for these specimens are available from me. Maps
represent distribution records from collections. For
some common and well known species (such as £.
bothrops, E. compositus, E. crotalinus, E. designatus), 1
include records from published sources I consider re-
liable.
Abbreviations for figure legends and synonymies
are: abd. = illustration(s) of abdomen; app. = illustra-
tions(s) of caudal appendages; cat. = listed in cata-
logue, descr. = description of male, female, larva, or
all, may include keys; distr. = documents or mentions
distribution of taxon; ept. = illustration(s) of epi-
proct; sep. = [page number of] separate (see Cowley
(1937) for pagination of reprint of Selysian mono-
graphs and synopses of Odonata); thx. = illustra-
tion(s) of thorax; vl. = illustration(s) of vulvar lami-
nae.
Abbreviations in descriptions are as follows: anx =
antenodal crossveins, pnx = postnodal crossveins, cs =
crossveins.
New names proposed in this paper follow the
ophidian example set by Selys, Hagen, Calvert, and
Williamson. All should be considered as nouns in ap-
position.
Most illustrations are from type material, which I
believe will aid further researchers when describing
new species. All illustrations were executed with the
aid of a camera lucida with a Wild M-8 stereoscopic
microscope.
Erpetogomphus Hagen in Selys, 1858
Erpetogomphus Selys, 1858 (Type species Ophiogomphus cro-
talinus [Hagen in Selys], 1854; designated by Kirby 1890:
61). — Selys 1858: 329 (69 sep.) (characters of genus);
Selys 1859: 535 (9 sep.) (addition of generic characters);
Hagen 1861: 98 (brief descr. of genus); Walsh 1863: 253
(comparison with other American gomphid genera);
Brauer 1868: 372 (in key to genera of Gomphidae);
Karsch 1890: 371 (mentioned in Gomphidae classifica-
tion); Calvert 1905: 147 (key to Middle American
Gomphidae); Calvert 1909: 468 (distr.); Calvert 1912a:
289 (distr. of genus); Kennedy 1917a: 544 (comments on
larvae); Tillyard 1917: 296 (distr.); Ris 1921: 343 (com-
parison with Mesogomphus [= Paragomphus]); Garman
1927: 125 (note on genus); Needham and Heywood
1929: 78 (descr.); Byers 1930: 52 (descr.); Cowley 1934:
241 (nomenclature of Erpetogomphus); Tinkham 1934:
218 (comments on distr. in Tex.); Fraser 1940: 544 (pos-
sible affinities with Onychogomphus based on penis mor-
phology); Needham 1941: 240 (comments on larva);
Needham 1944: 172 (verification table, larva); Wright
and Peterson 1944: 152 (in key to larvae of nearctic
Gomphidae); Needham & Westfall 1955: 139 (treat-
ment of genus); Pritchard & Smith 1956: 114 (key to lar-
vae of nearctic Gomphidae, key to genera of
Gomphidae); Ferguson-Beatty 1956: 369 (proventriculus
of larva); Musser 1962: 13 (diagnosis of larvae); Gloyd
1963: 147 (molar structure of adult); Westfall 1984: 158
(in key to larvae of nearctic Gomphidae); Chao 1984: 79
186
(uncertainty of Ærpetogomphus in subfamily
Onychogomphinae); Carle 1986: 313 (characters of ge-
nus within Onychogomphini); Belle 1988: 99 (in key to
neotropical genera of Gomphidae); Bridges 1991: II.17
(cat); Novelo-G. & Pena-O. 1991: 129 (origin of
Erpetogomphus in Mexico); Gonzalez-S. & Novelo-G.
1991: 97 (adults in Mexico); Novelo-G. & Gonzalez-S.
1991: 150 (larvae in Mexico); Belle & Quintero 1992: 93
(in key to adults and larvae of Panamanian Gomphidae);
Carle 1992: 148 (key, Erpetocyclops subg. n. [type = E.
ophibolus Calvert], Calogomphus subg. n. [type = E. eutai-
nia Calvert]).
Herpetogomphus Walsh, 1862: 388 (unjustified emendation
of Erpetogomphus). — Hagen 1875a: 42 (cat.); Selys 1879:
63 (2 sep.) (modification of generic characters); Kirby
1890: 60 (cat.); Carpenter 1897: 452 (distr.); Needham
1897: 166 (in key to nearctic Gomphidae); Needham
1899: 234 (venation); Calvert 1899: 386 (comments on
Selys’ classification); Needham & Hart 1901: 53 (in key
to larvae of North American Gomphidae); Férster 1914:
73 (comparison with Ammogomphus); Seemann 1927: 19
(key to larvae, adults of western U. S. Gomphidae);
Needham 1940: 389 (key, verification table).
Medium sized to small neotropical Gomphidae of
slender build.
Head. — Eyes widely separated, characters of the
mouth parts, clypeus, frons typical of the
Gomphidae. Vertex variable, often interspecifically
and, in the £. ophibolus group, sexually dimorphic.
Anterior margin of vertex with a transverse trough, or
a pair of transverse pits anterior or anterolateral to
median ocellus, median ocellus anterior to lateral oc-
elli, or in some members of the E. ophibolus group,
this structure at posteriormost part of deep, longitu-
dinal trough, thus causing median ocellus to be pos-
terior to lateral ocelli (figs. 145-146); median surface
of vertex at vicinity of lateral ocelli and postoccipital
tubercle mostly planar, but members of £. ophibolus
group with a complete postocellar ridge, which may
be bilobed (E. agkistrodon, fig. 148) or, in remainder
of group, vestigial; occiput highly variable according
to species and sex, E. ophibolus group with occiput
small, semicircular, in other groups roughly trape-
zoidal, with posterior part broader than anterior part,
its dorsal surface roughly planar to nearly vertically
inclined, some species with slight to strongly tumid
area medially, one species (female of E. /eptophis) with
a posteriorly projected medial spine; rest of occiput
variable, semicircular, linear, weakly to strongly
notched medially, or sinuate; condition of crest often
variable within a few species; postocciput variable,
most in the £. ophibolus group convexly arcuate, in
others slightly convex to linear, postocciput mostly
not visible from above, but conspicuously so in others
(e.g., female of E. compositus); lateral margins of one
species (females of E. cophias) with transverse depres-
sions.
Thorax. — Prothorax and synthorax typical of the
Gomphidae, but with great differences in maculation
and colouration due to species and age differences.
Pale colour pale green, apple green to vivid blue
green, usually lighter ventrally and often with a slight
dusting of white pruinosity ventrally and around cox-
ae. Synthorax immaculate to heavily patterned with
full complement of thoracic stripes. Legs moderately
short, with femora slightly swollen and usually pale
basally, with dark brown to black distally; tibiae in al-
most all species black, prothoracic tibiae of males
with small keel occupying distal 0.15 of mesal ventral
margin.
Wings (figs. 227-230) variable inter- and intraspe-
cifically. Fore wing moderately narrow, with 11-21
antenodal crossveins, 5-16 postnodal crossveins, no
subcostal crossvein; pterostigma prominent, about 4
times as long as wide, widest medially with its poste-
rior margin slightly convex, surmounting 4-8 cross-
veins, the proximal a brace vein; anterior margin of
costa in many species pale, becoming dark at pteros-
tigma; arculus at vicinity of second antenodal, the
sectors not stalked, separated at base by distance
greater than width of each sector; supratriangular, tri-
angular, and subtriangular crossveins absent; paranal
cells variable in number, often numbering 5-7, small-
er species usually with zero or few supplementary
marginal cells behind paranal cells, other species with
irregular row of up to 6-7 supplementary cells; no ap-
ical or medial planates, discoidal field of two rows.
Hind wing with 7-14 antenodal crossveins, 7-14
postnodal crossveins; anal triangle of 2-4 cells, often
with one of them in the latter condition small; anal
loop absent (some specimens with a vestigial anal
loop of 3 cells, but this condition rare); one paranal
cell at Y interspace, two paranal cells at X interspace.
Wing colouration entirely hyaline or flavescent at
bases in some species. Hind wing length 21-36 mm.
Abdomen cylindrical, narrowest medially in males,
transverse carinae on segments 3-7; auricles well de-
veloped in male, inner margin denticulate; auricle
vestigial in female; male posterior margin of tergites 8
and especially 9 in a few species prolonged into a me-
dial point, but this condition variable intraspecifical-
ly; segments 8-9 moderately clubbed, with moderate
foliate extensions; female with apical abdominal seg-
ments hardly widened, foliar expansions small.
Colour pattern and maculation variable between spe-
cies and within sexes; all species with a pale green to
pale orange middorsal stripe, sometimes extending
full length of segment, but progressively less so on
more posterior segments; dark brown to black dorso-
lateral stripe present on first 7 segments, usually con-
stricted or separated medially, giving most of abdo-
men a serial dark and pale appearance; posterior part
of segment 7, all of segments 8-10 tan to dark red
brown, especially middorsally and ventrolaterally.
GARRISON: Revision of Erpetogomphus
Length of abdomen including appendages: d : 29-40
mm, 9: 29-41 mm.
Abdominal appendages of male with cerci of great
diversity according to species, but these structures al-
ways as long as segment 10, cercus usually gently de-
cumbent at apical 0.30; rarely with a dorsal append-
age. Ventral carina present basally, or distally, or
absent, depending on species, epiprocts divided at
base but contiguous along most of their length, gent-
ly to strongly curved at distal 0.50 to 0.30, depending
on species; this structure lacking any accessory spines,
or protuberances (except for E. ophibolus, fig. 95); tips
of epiprocts assuming a wide variety of shapes and
forms, and often variable intraspecifically (e.g, E.
bothrops, figs. 127-128).
Vulvar lamina of female variable interspecifically,
but never more than 0.50 length of segment 9, vulvar
plates small, digit-like to forming large, contiguous,
triangular or quadrangular plates, a moderate to large
V- or U-shaped notch medially; postlamellar ridge al-
ways present, semicircular in only two species (E. eu-
tainia, E. leptophis), in all others forming an inverted
Y; stem of Y originating before or often at level of vul-
var plates, some species with a well defined semicircu-
lar depression on each side of posterior stem.
Male genitalia variable among species: anterior
lamina not prominent; anterior hamule smaller than
posterior hamule, usually with the distal end hidden
from view by posterior lamina; anterior lamina dark,
divided at various heights according to species, but
with posterior branch usually less conspicuous, small-
er, and anterior branch more prominent; posterior
lamina usually pale, linear in two species (E. eutainia,
E. leptophis); in all others with a moderately wider
base, its tip usually adorned with an apical or anteap-
ical hook directed cephalad or cephalolaterad. Penis
with basal (first) segment prominent, with a semicir-
cular rim, especially so laterally, posteriorly with a
pair of prominent lobes, or these lobes vestigial; sec-
ond segment moderately long, third short, with a pair
of small, nipple-like sclerotizations dorsally in two
species (E. eutainia, E. leptophis); fourth segment
highly variable specifically, with or without a pre-
puce, most species with a well defined lateral lobe; in
most of the £. ophibolus group, lateral lobe small and
pointed; membranous hood short to long and over-
lapping; a pair of flattened membranous cornuae of
various lengths and shapes present on apical part of
segment; in most of the E. ophibolus group, cornuae
absent or vestigial; penis guard short, quadrate in
cross-section.
A more complete discussion of the male and female
genitalia is given under discussions of the species
group accounts and in the cladistics section.
187
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Keys
The keys should work for most specimens, but
the worker may have difficulty when trying to deter-
mine poorly preserved material. I have used morp-
hological characters for both sexes wherever possi-
ble, but some species pairs (eg, females of E.
heterodon and E. sipedon, or E. tristani and E. sabale-
ticus) show little or no morphological differentia- ‘
tion. For them, I have had to rely on colour pattern.
Abdomens of some females, particularly those late-
rally compressed, may need to be relaxed so that the
vulvar lamina can be seen.
= to Erpetogomphus males
DD)
4 (3).
5 (1).
6 (5).
Cercus with a prominent superior tooth at
about 0.75 of appendage length (figs. 92-94)
Cercus with dorsal surface convexly angulate
(fig. 112), smoothly curved (fig. 103),
straight (fig. 100), or with a concavity (fig.
IS) ete Me eea EE 5
Epiproct smoothly curved, with tip pointing
dorsally or posterodorsally, a small, sharp,
anteriorly directed superior tooth at middle
of epiproct (fig. 95). Southern Mexico,
Belize Mana dieu na fedi; du ophibolus
Epiproct strongly curved so that distal 0.30
is parallel to basal 0.30, tip of epiproct
pointing anteriorly, superior surface of epi-
Proc@with noktoother en 3)
Ventral margin of cercus smoothly concave
(fig. 92). Northeastern Mexico to Costa
Rica. a ie ica ees cee) RA constrictor
Ventral margin with a large ventral tubercle
at 0.50-0.75 length of cercus (figs. 93-94).
Costa Rica south to Colombia and
Venezuela she. ti ende im, 4
In lateral view, superior tooth of cercus as
long as rest of appendage, so that appendage
appears to end in two equal branches
(fig. 93). Panama, Colombia, Venezuela .....
elen Sods tle Seti topo Siae caldi: sabaleticus
In lateral view, superior tooth of cercus less
than 0.50 as long as remainder of appendage
(fig. 94). Costa Rica, northern Panama
ee RE aa e de tristani
Metathoracic tibiae either entirely yellow ex-
ternally (fig. 169), or yellow with median
longitudinal black line. Western Mexico,
southeastern Arizona ................... crotalinus
Metathoracic tibiae entirely brown or black
externally; never with any yellow (e.g, figs.
168211 0) een 6
Basal 0.25 to 0.30 of lower margin of cercus
with a distinct longitudinal carina which
183
7 (6).
11 (10).
12 (10).
may end in a small ventral tooth (e.g., E. co-
iphiaspfigal oO). EN 7
Basal part of cercus with no ventral carina,
though a small tubercle may be present at
basal 0.15-0.20 of appendage (e.g., figs. 99,
OD Seti IE Soa ORTE en 14
With a dark stripe bordering posterior mar-
gin of metepimeron (figs. 8-9). Southern
Mexico, Belize through Costa Rica .............
geler Aspirine Aen PA eutainia (in part)
Without a dark stripe on posterior margin of
Metepimeron … avro vennen 8
In lateral view, ventral margin of cercus with
carina terminating in a distinct ventral tooth
at basal 0.30 of appendage (fig. 116); thorax
almost immaculate, at most barely an indi-
cation of a dark antehumeral stripe (fig. 27).
Michoacan and Morelos states of western
Mericons, acuta are cee cophias
Ventral margin of cercus with longitudinal
carina not terminating in a ventral tooth;
thorax with well-defined dark antehumeral
stripe, though it may be an elongated, isolat-
edispoti(erg figs LI 9
Dorsal surface of cercus distinctly angulate
invlateralaviews (Gios 100} 102) WA 10
Dorsal surface of cercus smoothly curved
(enden anice 13
Sides of thorax almost entirely green, with
only a small, ill-defined dark humeral stripe;
second lateral stripe incomplete, dark only
below metathoracic spiracle, or absent; and
third lateral stripe present only on upper
0.30 of suture (fig. 23), or absent (fig. 19)...
Sides of thorax green with usual comple-
ment of dark thoracic stripes; second lateral
stripe may be incomplete, but humeral and
third lateral stripes always present (figs. 17,
2020) wee ss, EL RE 12
Dark antehumeral stripe incomplete, not
touching humeral stripe (fig. 23); tip of cer-
cus acute, but not gradually narrowing pos-
teriorly (figs. 110-111); cornuae of penis
without mesal lobes (fig. 87). New Mexico,
western Texas, northern Mexico .................
Dark antehumeral stripe complete, usually
connected to humeral stripe (fig. 19); tip of
cercus drawn out into a narrow cylindrical
point (figs. 105-106); cornuae with mesal
lobes (fig. 83). Durango, Jalisco, Morelos,
and Puebla states of central Mexico ............
vn eeN Fei leer sipedon
Tip of cercus strongly acuminate (fig. 104),
dark antehumeral stripe not connected to
1510)
14 (6).
17 (16).
collar (fig. 17) (except in some Mexican in-
dividuals), base of wings flavescent (except
in some Mexican individuals, fig. 18), me-
dian area of occiput with a strongly raised
tubercle (fig. 156). Eastern U.S. to Arizona,
northern Mexico … designatus
Tip of cercus blunt, not acuminate (figs.
107-108); dark antehumeral stripe always
connected to collar (ssp. matrix, fig. 21), or
largely connected to humeral stripe (ssp.
lampropeltis, fig. 20); base of wings hyaline,
median area of occiput only slightly raised
(fig. 160). Southern California, Arizona,
New Mexico, western Texas, Baja
California, Durango state, Mexico ..............
mito Lecter ET TEE lampropeltis
Ventral carina at base of cercus black, con-
trasting with remainder of appendage; this
carina usually denticulate; thorax blue-green
in life; sides of posterior hamules parallel,
linear (fig. 52). Southern Texas to southern
Mexico es eutainia (in part)
Ventral carina at base of cercus pale, the
same colour as rest of appendage, this carina
smooth; thorax pale green in life; sides of
posterior hamules converging toward tip,
not linear (fig. 66). Southwestern United
States BajalCalitorniar ento compositus
Distal 0.5 of cercus concave dorsally, this ar-
ea covered with thick, long bristles (fig.
115). Southern Veracruz state, Mexico … …
Distal 0.5 of cercus straight or convex (e.g.,
figs. 99, 103)
. Ventral margin of cercus straight (figs. 99,
100) or forming a gentle concave curve (fig.
98) so that appendage appears linear ..... 16
Apical 0.30 of ventral margin of cercus
Clnvedid ow WA eneen tease cece 18
. Thorax entirely yellow-green (fig. 13), or
with only an antehumeral and faint indica-
tion of a dark middorsal and antehumeral
stripe (figs. 11-12); face entirely pale.
MEsicosonth to CostaRies mr. 17
Thorax blue-green with full complement of
dark thoracic stripes (fig. 10); face blue-
green with extensive dark brown frontocly-
Pealestnipem RO nee nn leptophis
Distal branch of anterior hamule thick, as
high as gap separating the two branches (fig.
54); posterior hamules with no distal tooth
(fig. 54). Guatemala south to Costa Rica ....
RA A MN sen len dist as elaphe
Distal branch of anterior hamule narrow,
not as high as gap separating the two
branches (figs. 55-56); posterior hamules
18(15).
19(18).
0 (19).
GARRISON: Revision of Erpetogomphus
with a distal tooth (fig. 55). Mexico to
Guatemala More ARR eee elaps
Thorax blue-green, with full complement of
dark stripes, including second lateral stripe
and stripe on posterior margin of metepime-
ron (fig. 6); face (fig. 142) predominantly
dark, with a well defined blue green spot on
frons. Guatemala, Costa Rica ......... schausi
Thorax blue-green or yellow-green; sides
lacking complete dark second lateral stripe
(except in some viperinus incomplete in
some bothrops) and without dark stripe on
posterior margin of metepimeron; face pre-
dominantly pale, with dark areas limited to
suture ei asa Rs a Re no 19
Tip of epiproct in posterior view terminat-
ing in a blunt point (fig. 129); southern
Veracruz state, Mexico................... viperinus
Tip of epiproct in posterior view spatulate
(fig. 126), broadly truncate (fig. 127), or bi-
dentate 23) e 20
Larger species (hind wing 30-31 mm); epi-
procts in lateral view curved, extending 0.50
or less the length of cerci (fig. 101), epi-
procts in posterior view truncate or slightly
bidentate; dark antehumeral and humeral
stripes combined or nearly so (fig. 14); pos-
terior hamular tooth prominent, directed la-
terocephalad or cephalad. Nuevo Leon,
Michoacan, San Luis Potosi states, Mexico.
(herl) iena mener liopeltis
Smaller species (hind wing 21-28 mm); epi-
procts in lateral view curved, extending
0.75- 0.90 the length of cerci (fig. 102), epi-
procts in posterior view bidentate or trun-
cate (fig. 128); dark antehumeral and hu-
meral stripes separate (fig. 15); posterior
hamular tooth not as prominent, more
strongly directed cephalad. Tamaulipas,
Mexico, south to El Salvador (fig. 199)
potent | nth beat he anca vele bothrops
o to Erpetogomphus females
Medial area of occiput with a posteriorly
pointed protuberance (fig. 151) … leptophis
Medial area of occiput variously dhaned, but
never with a posteriorly directed protube-
rance
Occiput with a shallow oval depression on
each side posterior to elevated rim (fig. 165).
Michoacán and Morelos, Mexico … cophias
Occipital area posterior to elevated rim en-
tire, with no shallow depressions
Outer surfaces of metathoracic tibiae entire-
ly yellow, or yellow with a median longitudi-
nal black line (fig. 169). Western Mexico,
189
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
10 (9).
southeastern Arizona ................... crotalinus
Outer surfaces of metathoracic tibiae entire-
ly black (e.g., figs. 168, 170)
Median ocellus posterior to lateral ocelli, the
former lying within a deep longitudinal
trough (figs. 145-146)
Median ocellus at level of or anterior to lat-
eral ocelli, no longitudinal trough ............ 6
Dorsal surface of vertex with sides of trough .
roughly parallel, or only slightly converging
anteriorly (fig. 145). Northeastern Mexico,
SOuUtMtO\ Costa Rica peewee sees constrictor
Dorsal surface of vertex with sides of trough
strongly converging anteriorly ......(fig. 146)
Costa Rica, western Panama ........... tristani
Central Panama, Colombia, Venezuela ......
AA OL Et MEE pi DUI sabaleticus
Vertex with transverse ocellar ridge bilobed
behind median ocellus (figs. 148-149) ..... 7
Vertex with transverse ocellar ridge entire
(fig. 147), low and almost vestigial (fig.
150), or absent, its lateral ends forming oval
tubercles posterior to lateral ocelli (e.g., fig.
150)
Occiput transversally narrow, forming a
shallow semicircle (fig. 148). Jalapa, Mexico
Erten, Sau Wes. hamteleren er agkistrodon
Occiput wide, forming a full semicircle (fig.
149). Guatemala, Costa Rica .......... schausi
Vertex with transverse ocellar ridge entire,
prominent, and concave posteriorly (fig.
147). Southern Veracruz, Mexico, Belize …
D PE SIATE LITRO à UE. ATARI ophibolus
Vertex with median area bare, or at most on-
ly slightly raised (e.g., fig. 150)
Vulvar lamina followed on segment 9 by dis-
tinct and prominent semicircular ridge, nev-
er with a posteriorly directed arm (fig. 178).
Southern Texas, eastern Mexico, Belize to
Guatemala and Costa Rica ............ eutainia
Vulvar lamina followed on segment 9 by a
Y-shaped ridge (e.g., fig. 195) ................ 10
Vulvar lamina with each plate strongly cor-
rugated diagonally, distal and lateral mar-
gins of each plate forming a strongly raised
semicircular ridge (figs. 195-196); a median
tumid swelling on postocciput immediately
behind occipital ridge (fig. 167) [includes %-
opeltis, but properly goes to 10’]. Southern
WeractuzN lexicon ee boa
Vulvar lamina with each plate entire (fig.
185) or bisected diagonally by only one
groove (fig. 186); distal and lateral margins
of each plate planar, not forming a strongly
raised semicircular ridge; postocciput not
differentiated, evenly curved (fig. 166) [tu-
190
11 (10).
12 (11).
3202)!
14 (13).
Sd
16105):
nt LINE De DE 11
Median surface of occiput with a strongly
raised tubercle (fig. 156). Eastern United
States west to Arizona, northern Mexico .....
A LE OE UE DS (IM DATE designatus
Median surface of occiput planar or at most
slighitlyeraisecs (108152) 12
Lobes of vulvar lamina separated by an al-
most U-shaped interval 3 to 4 times as wide
as either lobe (figs. 180-181)
Mexico south to Guatemala
Guatemala to Costa Rica .................. elaphe
Lobes of vulvar lamina separated by a trian-
gular or semicircular interval 0.5 to 1.0 the
width of each lobe (e.g., figs. 182, 187) ...13
With a second complete dark lateral stripe
on thorax (e.g., figs. 20-21, 24) … 14
Second dark lateral stripe completely lacking
or vestigial, at most extending from base to
just above metathoracic spiracle (e.g, fig.
36)
Occiput in dorsal view narrow, its width less
than width between median ocellus and oc-
ciput; postoccipital area easily visible (fig.
163); base of wings with flavescent infusion
between Sc and R at least up to first anx and
cubitoanal area; top of abdomen with light
areas ivory or with a tinge of orange, usually
not different from white on sides.
Southwestern United States, Baja California
AT RA ARIA E LO SIA compositus
Occiput in dorsal view wide, its width al-
most equal to width between median ocellus
and occiput; postoccipital area not visible
(fig. 160); base of wings hyaline; top of ab-
domen with light areas much darker than
white on sides. Southwestern United States,
Baja California, Durango state, Mexico ......
N ec PISA lampropeltis
Occiput narrow, vestigial, with only a small
medially arched area (fig. 153); post-occiput
with a median tumid area (as in fig. 167).
Nuevo Leon, Michoacan, San Luis Potosi
STALES IMIEXICO ME TE II liopeltis
Occiput not especially narrow, extending
full width (e.g. figs. 154-155, 159, 162);
post-occiput planar or, at most, only slightly
convex medially (as in fig. 166)
Stem of postlamellar ridge with a well de-
fined circular or oval depression on either
side; junction of Y of postlamellar ridge usu-
ally posterior to hind margin of vulvar lami-
nanplates (METEO E 17
Stem of postlamellar ridge with only a slight
depression or, more often, without any cir-
cular or oval depression on either side; junc-
17 (16).
18 (16).
tion of Y of postlamellar ridge at or anterior
to hind margin of vulvar laminar plates (figs.
SENS SMS) aoe e ne 18
A deep pit at anterior margin of frons ante-
rolateral to median ocellus (fig. 155); cleft
between vulvar laminar plates wide, forming
an obtuse arc greater than 100°; dark hu-
meral and antehumeral stripes combined or
nearly so (fig. 16); larger species (hind wing
31-34 mm). Southern Veracruz, Mexico ....
OBS SSA A DBR OAM ha Hae A SDE Un LET viperinus
Anterior margin immediately anterior to
median ocellus forming a narrow V-shaped
trough with base of postfrons (fig. 154); cleft
between vulvar laminar plates narrow, form-
ing a V- shaped notch of not more than 90°
(fig. 183); dark humeral and antehumeral
stripes separate (fig. 15); smaller species
(hind wing 17-31 mm). Tamaulipas,
Mexico, south through El Salvador ............
Stic BORSA NRE OES ERE TER bothrops
Dark antehumeral stripe complete, usually
connected at its upper end to narrow hu-
meral stripe (fig. 19). Jalisco, Morelos, and
RueblawVicxicon eee ae sipedon
Dark antehumeral stripe vestigial, forming
an isolated elongate spot, not connected to
vestigial humeral stripe (fig. 23).
Southwestern United States, northern
Mexico Kebe a ei heterodon
Key to Erpetogomphus males based primarily on
characters of abdominal segments 2 and 3
I include this key for male specimens lacking the
posterior abdominal segments. The key should allow
a high probability of identification, though some spe-
cies (E. ophibolus, E. sabaleticus, E. tristani, E. constric-
tor, for example) appear inseparable by penis struc-
ture and are distinguished only by the caudal
appendages. For those species, I have added other so-
matic characters and/or geographic data with which
to separate them. The key does not include E. agkis-
trodon, the male of which is unknown.
Ik
Ental surface of penis with a prepuce (e.g.,
fig. 76), this structure reduced and partially
hidden by frill-like lateral lobe in leptophis
(fig. 76) and eutainia (fig. 75); thoracic pale
colouration in life deep blue green, dark tho-
racic stripes always present on sides (e.g., fig.
UO), Rene SEGRE a es, N
Ental surface of penis lacking a prepuce (e.g.,
fig. 90); thoracic pale colour in life yellow
green (except for viperinus and probably lio-
peltis); dark lateral thoracic stripes often (but
not always — figs. 16, 18) reduced or absent
GARRISON: Revision of Erpetogomphus
Posterior hamule digit-shaped (figs. 52-53),
its tip broadly rounded; anterior hamule di-
vided at upper 0.30 to 0.25, its posterior
branch well defined, its tip almost meeting
larger anterior branch (figs. 52-53); lateral
lobe of penis composed of large spinulose
frill (figs. 52-53); cornuae well developed,
theinfapicesipointed ee 3
Posterior hamule triangular, its tip armed
with a spine (fig. 47); anterior hamule with
only a posterior shoulder (e.g., fig. 47); later-
al lobe of penis reduced to a small, semicir-
cular, spinulose frill (fig. 74) or vestigial (fig.
70); cornuae vestigial (fig. 74) or present
(Hg 70) EERE ENEN 4
Membranous hood of penis about 3 times as
long as wide, apices overlapping (fig. 76)
[Note: this character is based on the only
known specimen, the holotype; other speci-
mens may show variability similar to eutai-
nia.|; dark second and third lateral stripes
connected at upper ends, forming an isolat-
ed pale spot below subalar carina; dark later-
al stripe along posterior margin of metepim-
eron well defined (fig. 10); caudal
appendages as in fig. 98; Belize ..... leptophis
Membranous hood of penis reduced, less
than 2 times as long as wide, though apices
may overlap (fig. 75); dark second and third
lateral thoracic stripes not connecting (fig.
7-9); dark lateral stripe along posterior mar-
gin of metepimeron absent (fig. 7) or
present primarily in populations of central
Mexico (fig. 8) south through Costa Rica
(fig. 9); caudal appendages as in fig. 97 ......
PARE REE cor Eee TEE re eutainia
Cornuae of penis present; their tips diver-
gent (fig. 74); lateral lobe a small, circular,
spinulose frill (fig. 74); Guatemala and
Costa RICA EE a Ae ent schausi
Cornuae vestigial (fig. 71); lateral lobe vesti-
gial, forming a small, acute point (fig. 71) ..
Appendages as in fig. 93; Central Panama
south to Colombia and Venezuela
LR i MORAR eee sabaleticus
Mexico south through Costa Rica and
MOTO AMATA nni ILI IONI NOEIRINIIIINIG0INO 6
Epiproct smoothly curved, with tip pointing
dorsally or posterodorsally, a small, sharp,
anteriorly directed superior tooth at middle
of epiproct (fig. 95). Southern Mexico,
Belize tota ee ts ophibolus
Epiproct strongly curved so that distal 0.30
is parallel to basal 0.30, tip of epiproct
191
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
BMD
pointing anteriorly, superior surface of epi-
proct with no tooth
Appendages as in fig. 92; Mexico south
through Costa Rica constrictor
Appendages as in fig. 94; northern Costa
Rica through northern Panama tristani
Anterior hamule divided at base; its branch-
es resembling a claw; posterior hamule
broadly spatulate (figs. 68-69)
Anterior hamule divided at lower 0.30 to’
upper 0.25 posterior arm short and truncate
or acute; posterior hamule broadly triangu-
ge: antenne aut hi eit
Dark antehumeral and humeral stripes unit-
ed, forming a broad, dark stripe over first
lateral suture (fig. 26); southern Veracruz,
México den helene clara boa
Thorax almost entirely pale, with only a ves-
tigial antehumeral stripe (fig. 27);
Michoacan and Morelos states of Mexico ...
CRE a BAS tity bee argent MAME A cophias
Lateral lobe of penis linear, at least 3 times
longer than wide; its posterior margin spinu-
lose (Cts, 777.15 82, Ge) eme 11
Lateral lobe of penis round, about as long as
wide, its margin spinulose (fig. 86) 16
. Lateral lobe of penis almost recumbent
against ental margin of fourth segment (fig.
82); anterior hamule divided at distal 0.5 of
segment; posterior hamule lacking a distal
tooth (fig. 60); posterior margin of first seg-
ment (peduncle) of penis strongly cleft, each
lobe on either side of cleft protruding well
beyond posterior margin of segment; south-
ern United States west to Arizona, northern
Mexico designatus
Lateral lobe of penis prominent, directed al-
most perpendicularly to ental margin of
fourth segment; anterior hamule divided at
distal 0.30 of segment, posterior hamule
with a distal tooth (except for elaphe from
Guatemala south through Costa Rica); pos-
terior margin of first segment of penis weak-
ly bilobed, lobes on either side of emargina-
tion only slightly extending beyond
posterior margin of segment; northern
Mexico south through Costa Rica ......... 12
Distal branch of anterior hamule thick, as
tall (high) as gap separating the two branch-
es (fig. 54); posterior hamule with no distal
tooth (fig. 54); Guatemala south to Costa
Rica elaphe
Distal branch of anterior hamule narrow,
not as tall (high) as gap separating the two
branches (figs. 55-56); posterior hamule
with a distal tooth 13
192
502).
14 (13).
15 (14).
16 (10).
Distal tooth of posterior hamule bent anteri-
orly so that it lies in same direction as the
planar surface of hamule (figs. 55-56);
Mexico south to Guatemala ................ elaps
Distal tooth of posterior hamule bent later-
ally or anterolaterally so that it points distal-
ly and perpendicularly from planar surface
of hamule (fig. 59) [Note: This condition
appears universally only in E. viperinus, but
variation in this condition occurs in E. both-
rops and E. liopeltis, some specimens of these
last two species may not be separable from
E. elaps by this character]; Mexico south
throuehi@uatemala er re ee
Base of postfrons medially with a pair of
deep pits, each anterolateral to median ocel-
lus (fig. 137). Southern Veracruz, Mexico ..
RR EN ade annen viperinus
Base of postfrons medially with a transverse
arcuate trough immediately anterior to me-
dian ocellus (fig. 136) 15
Smaller species (hind wing 21-28 mm); dark
antehumeral and humeral stripes separate
(fig. 15); posterior hamule with apical tooth
shorter, less prominent, more strongly
curved laterally over tip of hamule.
Tamaulipas, Mexico, south through El
Salvador (fig. 199) bothrops
Larger species (hind wing 30-31 mm); dark
antehumeral and humeral stripes connected
(fig. 14); posterior hamule with apical tooth
longer, more linear, not as strongly curved
laterally over tip of hamule. Nuevo Leon,
Michoacan, San Luis Potosi, Mexico
liopeltis
Cornuae of penis with tips long, narrow,
and attenuate; medial lobe present (fig. 86);
extensor surfaces of tibiae yellow, or yellow
with a median longitudinal black line;
south-eastern Arizona south through central
Mexico crotalinus
Cornuae of penis with tips bluntly rounded
(figs. 87-88); medial lobe present (figs. 83-
84) or absent (figs. 87-88); extensor surfaces
of tibiae black, never with yellow … … …
Tips of cornuae bluntly rounded; medial
lobe absent (figs. 87-88) 18
Tips of cornuae with a well defined medial
lobe (figs. 83-84)
Median part of base of postfrons with a
transverse arcuate trough immediately ante-
rior to median ocellus (as in fig. 138); west-
ern United States compositus
Base of postfrons medially with a pair of
deep pits, each anterolateral to median ocel-
lus (as in fig. 138); western Texas, western
19 (17). Penis in dorsal view with membranous hood
short, exposing a subcutaneous membra-
nous hood which forms the posterior margin
of segment [be sure to examine closely: rota-
tion of penis to dorsolateral view will reveal
a raised hood anterior to subcutaneous
membrane], (fig. 83); Jalisco, Morelos, and
Puebla, Mexico ............ sipedon
= Penis in dorsal view with membranous hood
long, completely covering subcutaneous
membrane, thus forming posterior margin
of segment (figs. 84-85); western Texas,
southern and central New Mexico and
Arizona, southwestern California, Baja
California, Durango state, Mexico ..............
ERE ENG BO sie ees lampropeltis
Erpetogomphus constrictor Ris
(figs. 1-thx, 28-abd, 97-hamules, 70-penis, 92-app,
145-® vertex, 171-vl, 198-distr)
Erpetogomphus constrictor Ris, 1917: 154 (Nach 3d, 19
von Misantla, Vera Cruz, Mexiko, VI 1911, die ich dem
schweizerischen Sammler W. Gugelmann verdanke.’). —
Williamson and Williamson 1930: 12 (summary of stat-
us); Montgomery 1973: 239 (derivation of name);
Paulson 1982: 255 (Mex.); Davies and Tobin 1985: 27
(cat.); Tsuda 1986: 87 (cat.); Dunkle 1988: 46
(Honduras); Maes et al. 1988: 36 (Nicaragua); Bridges
1991: VII.51 (cat.); Tsuda 1991: 95 (cat.).
Description
Male. — Labium grey, becoming dark grey medial-
ly, labrum entirely brown to grey green with dark
brown margin and large inverted medial triangular
spot; base of mandibles grey green, anteclypeus grey
green; postclypeus brown except for small triangular
green spots above lateral lobes; ventral margin of
frons brown joining brown of postclypeus, thus for-
ming a large brown frontoclypeal stripe; remainder of
frons green; extreme base of frons, vertex, and occiput
dark brown; area immediately anterior to median
ocellus with a shallow longitudinal trough, postocel-
lar tumid areas prominent, a lower incomplete trans-
verse ridge posterior to median ocellus but not conti-
nuous with each postocellar tumid area laterally;
occiput planar, its crest costate, linear, fringed with
dark brown hairs; postocciput brown, linear to gently
concave medially; rear of head brown.
Prothorax brown, paler dorsally, anterior and pos-
terior lobes green, some specimens with two small
green midlateral spots (almost touching) on median
lobes; synthorax (fig. 1) with prominent, well defined
dark stripes on green background as follows: middor-
sal stripe narrowing anteriorly at collar, costal mar-
GARRISON: Revision of Erpetogomphus
gins of antealar sinus and area along this structure
connecting with humeral stripe; antehumeral stripe
connecting with humeral basally and dorsally, but its
upper end often not touching brown below antealar
crest, or, if so, then isolating small area of green; sec-
ond and third lateral stripes connecting at upper
0.25, isolating small green spot, second lateral stripe
swollen in vicinity of metaspiracle and, in some spec-
imens, enlarged and touching third lateral stripe; pos-
terior margin of metepimeron, metasternum, and
venter of thorax grey green to tawny. Venter of profe-
mora grey green, remainder of femora dark brown,
becoming black distally; tibiae, tarsi, armature black.
Wings hyaline, venation and pterostigma black.
Venational statistics. Fifth (rarely fourth or sixth)
antenodal thickened; number of marginal cells be-
hind fore wing paranal cells: 0-1/0-1; anx: fore wing
13-17/14-17, hind wing 10-12/10-13; pnx: fore wing
10-14/9-14, hind wing 9-12/9-13; cs under pteros-
tigma.: fore wing 5-7/4-7, hind wing 4-7/5-7; anal
triangular cells: 3- 5/3-4. Hind wing 25-30 mm.
Abdomen with segment 1 brown, green laterally
and with a green middorsal stripe; segment 2 brown
except for green auricles, ventral margin, and narrow
middorsal stripe, annulus black; segment 3 primarily
black with green middorsal stripe extending from
black annulus of segment 2 and ending at distal annu-
lus, a pale grey green anterolateral spot becoming nar-
rower along ventral margin and disappearing at 0.25
to 0.75 of segment length; segments 4-6 similar to
segment 3 but with pale middorsal stripe ending at
posterior 0.10 to 0.25 of segment, segment 6 in some
specimens with anterolateral white connecting dorsal-
ly with pale green middorsal stripe; segment 7 grey
green on anterior 0.50 to 0.60, except for dark trans-
verse carina, posterior part of segment brown to
black; segments 8-10 red brown, becoming darker
dorsally and ventrally, especially along foliate mar-
gins, posterior medial margin of segment 9 bluntly
pointed or slightly so. Abdomen 31-35 mm.
Cercus (fig. 92) grey green, becoming dark brown
posteriorly, arcuate with a prominent laterally com-
pressed dorsal tubercle on distal 0.75 of cercus, ex-
treme base with a black ventral tooth (mostly hidden
by posterior margin of abdominal tergite 10), a ven-
tral carina on distal 0.25 of cercus, tip of cercus with
a black tooth; epiprocts dark brown, strongly curved
at distal 0.50 so that distal 0.5 of appendage is par-
allel, distal 0.30 sulcate medially on exterior surface.
Accessory genitalia. Hamules (fig. 47) small; anterior
hamule brown, divided at distal 0.30, posterior
branch a well developed shoulder; posterior hamule
roughly triangular, its tip with a spine; penis (fig. 70)
with cornuae slightly bilobate or transverse at base, a
well developed prepuce, lateral lobe a small tooth.
Female. — Head markings similar to male but with
193
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
dark median spot on labrum larger, often connecting
with ventral brown, thus separating green into two
lateral spots; some specimens with an entirely brown
labrum; brown on frontoclypeal suture more exten-
sive laterally, so that entire postclypeus may be
brown; brown at base of antefrons with a median ex-
tension which, in some specimens, connects with
brown frontoclypeal suture, thus dividing frontal
green into two spots. Morphology of vertex and occi-
put differs from male as follows: a large longitudinal
trough on vertex with median ocellus recessed poste-
rior to lateral ocelli (fig. 145); bottom of median fur-
row with longitudinal convex area; lateral margins of
trough formed by median ocellar protuberances con-
necting posteriorly forming a semicircular ridge; OCCI-
put small, forming a narrow semicircle, crest covered
with long brown hairs; postocciput brown, convex.
Pro- and synthorax as in male, but with dark areas
more extensive.
Venational statistics (n = 15). Fifth antenodal
thickened in all wings; number of marginal cells be-
hind fore wing paranal cells: 0-5/0-3; anx: fore wing
14-17/14-17, hind wing 11-12/11-12; pnx: fore wing
11-14/11-14, hind wing 10-14/11-13; cs under pte-
rostigma.: fore wing 5-7/5-7, hind wing 5-7/5-7.
Hind wing 28-31 mm.
Abdomen (fig. 28) with segment 1 brown, a green
spot posterolaterally and with a complete green mid-
dorsal stripe; segment 2 brown except for following
green areas: wide middorsal stripe narrowing posteri-
orly, longitudinal lateral stripe connecting just anteri-
or to auricle and narrowing slightly to posterior mar-
gin of segment, a thin pale margin at venter of
segment; segment 3 similar to segment 2 but lateral
pale stripe ending posteriorly at transverse carina,
middorsal green narrowly acuminate posteriorly,
forming a mere line on posterior 0.25 of segment;
segments 4-6 similar to segment 3 but with middor-
sal green progressively shorter posteriorly, lateral pale
stripe white and forming a small quadrangular spot at
anteroventral margin of segment; segment 7 with an-
terior 0.50 pale green, interrupted by black transverse
carina, posterior 0.50 becoming red brown; segments
8-10 all red brown, often darker dorsoposteriorly;
cerci pale grey green. Vulvar lamina (fig. 171) with
plates broadly connected at basal 0.75, relatively un-
specialized, cleft obtusely V- or U-shaped, about as
wide as each lobe; postlamellar ridge with Y-juncture
posterior to plates, base of stem well defined and with
oval depression laterally. Abdomen 30-35 mm.
Diagnosis
Males of E. constrictor are easily separated from its
closest allies, E. tristani and E. sabaleticus, by the sha-
pe of the cercus. This structure is concave ventrally in
E. constrictor, not convex as in the other two species.
194
It superficially resembles Æ. schausi in body coloura-
tion and is diagnosed under that species.
Females seem to be separable from the same two
species by the width of the median trough of the ver-
tex. This structure is slightly divergent anteriorly in £.
constrictor (figs. 145), but is largely parallel in the oth-
er two species (fig. 146).
Remarks
Variation. — Venational details of lectotype male of
E. constrictor. No marginal cells behind fore wing pa-
ranal cells; anx: fore wing 15/14, hind wing 11/11;
pnx: fore wing 11/12, hind wing 12/13; cs under pte-
rostigma: fore wing 7/7, hind wing 7/6; number of
anal triangular cells: 4/3. Hind wing 29 mm.
Material I examined indicates that specimens from
more southerly parts of its range (Guatemala,
Honduras, Nicaragua) are more melanic than those
from eastern Mexico. For example, black markings
on the labrum, postclypeus, and thoracic striping are
generally more extensive, further reducing or isolating
green areas.
Two females, one from San Luis Potosi (El Salto
Falls) and the paralectotype are unusual in having 5/3
and 4/3 marginal cells behind fore wing paranal cells
in right and left fore wings, respectively. All other fe-
male specimens ranged from 0-1 marginal cells.
Biology. — E. González (in litt.) found this species
to be abundant at the Rio Huichihuayan. Males
perched on leaves, vines, and snags at the edge of the
river. Judging from specimens in collections, E. con-
strictor is the northern counterpart of E. tristani, and
appears to be common in certain areas.
Erpetogomphus constrictor appears to be a lowland
species of wet tropical forest habitats along the east
coast of Mexico south through Costa Rica
(Guanacaste Prov.: Miravalles, 26 April 1991 (C.
Esquivel), 38, A. Ramirez, in litt.). Altitudes range
from near sea level (70 m, San Luis Potosi, km 410)
to about 550 m (Nuevo Leon, entrance to Horsetail
Falls). Collection dates range from 26 April to 4
October. One specimen was collected in an ultravio-
let light trap.
Distribution (fig. 198). — This species occurs along
the eastern Mexican states of Nuevo Leon,
Tamaulipas, San Luis Potosí, Veracruz, south
through Guatemala, Honduras, and Nicaragua, to
Guanacaste Prov., Costa Rica. It is apparently parapa-
tric or allopatric with £. tristani in Costa Rica.
Material
Type data. — Lectotype d by present designation
with the following data in Ris’ hand: ‘15098’ in green
ink in an unknown hand, Erpetogomphus | n. sp. ó /
constrictor Ris 1917/ Misantla, Vera Cruz / Mexico
VI 1914 / W. Gugelmann. Two paralectotype males
and one female with same data. One male is labelled
‘15092’ in green ink, another male in poor condition
(head partially fragmented, abdomen broken in sever-
al places) is labelled ‘Type’ probably by Ris. The left
pair of wings is missing and probably served for figu-
re 6 in the original description. The female is labelled
‘15096’ in green ink. In sMF.
Other material (534, 159, including lectotype d and
paralectotypes). — Mexico: Nuevo Leon: Cola Caballo, SW
Monterrey, 6 July 1960 (W. B. Cutts), 12 (ummz); Rio
Ramos, 3 km S of Allende, 500 m, 7 Sept. 1963 (T. W.
Donnelly), 18 (rwe); south of Monterrey, Rio Elizondo,
19-20 June 1965 (O. S. Flint, Jr.), 65 (USNM, RWG); stream
at entrance to Horsetail Falls, 550 m, 25 June 1965 (D. R.
Paulson), 16 (pre); Tamaulipas: Gomez Farias, Rio Frio, 4
Oct. 1985 (F. Arias, L. Cervantes, M. Garcia), ‘colectado en
trampa U.V. en la noche’, 16 (rw); San Luis Potosi: El
Salto, 4 June 1967 (O. S. Flint, Jr.), 12 (usnm); Palitla, N
of Tamazunchale, 25 June 1965 (O. S. Flint, Jr.), 9d
(USNM, RWG); Huichihuayan, rte 85, km 399, 7 Aug. 1966
(O. S. Flint, Jr.), 5d (UsNM, Cc, TWD); stream crossing Hwy
85 (1000’); 6 mi N of Tamazunchale, 4 Sept. 1957 (G. H.
Beatty, III), 1d (rsca); km 410, ‘Sam Brown’ Hacienda,
250 ft, 27 Sept. 1938 (L. J. Lipovsky), 18 (ummz); Route
85, km 399, Huehuetlán, 26 June 1965 (O. S. Flint, Jr.),
48,19 (usnm); Rio Huichihuayan, Mpio de Huchuetlan,
100 m, 26 May 1987 (E. González-S, 26 (RWG); upstream
Cascadas Micos just S of town N of Ciudad Valles off Hwy
85, 27 June 1990 (J. S. Daigle), 16 (RwG); GUATEMALA:
Suchitepequez Dept., Cuyotenango, Finca San Rafael
Olimpo, 1700 ft, 10-20 June 1966 (O. S. Flint, Jr., M. A.
Ortiz-B.), 88 (USNM, RWG); HONDURAS: Comayagua Dept.,
Rancho Chiquito, 11.8 mi SE Villa de San Antonio, 2-3
Aug. 1967 (O. S. Flint, Jr., M. A. Ortiz-B.), 3d, 72 (usnm,
cc, RWG); Francisco Morazan Dept, 30 km E of
Tegucigalpa, 13 June 1982 (Fernandez, Garcia), 19 (swp);
[no date], (Carlos Jaramillo), 12 (swp); Er SALVADOR:
Ahuachapan Dept., Loma de Paya, Bosque El Impossible,
San Francisco Menendez, 24 July 1987 (A. Campus), 1d
(vx); 20 Aug. 1986 (V. Hellebuyck), 19 (vx) 4 Aug. 1987
(V. Hellebuyck), 19 (vx); NIcARAGUA: Chobales Dept., La
Flor, rte 7, km 159, 4 mi W of Acoyapajit, 29 July 1967 (O.
S. Flint, Jr., M. A. Ortiz- B.), 4d (usnm); Managua Dept.,
stream along Pan American Hwy, 7 km S of Nandaime, 21
June 1962 (T. W. Donnelly), 15 (rwp).
Erpetogomphus sabaleticus Williamson
(figs. 2-thx, 48-hamules, 71-penis, 93-app, 172-vl,
198-distr)
Erpetogomphus sabaleticus Williamson, 1918: 1 (descr. holo-
type d , allotype ®). — Williamson and Williamson 1930:
12 (summary of status); Paulson 1982: 255 (Panama,
South America); Davies and Tobin 1985: 28 (cat.);
Tsuda 1986: 87 (cat.); De Marmels 1990: 338 (Venez.);
Bridges 1991: VII.184 (cat.); Tsuda 1991: 95 (cat.); Belle
1992: 32 (descr. larva); Donnelly 1992: 85 (Panama);
Belle and Quintero 1992: 99 (key, Panama).
Description
A thorough comparative description of the holoty-
pe male and allotype female is given by Williamson
Garrison: Special codes for printers
(1918). I illustrate the thorax (fig. 2), caudal appen-
dages (fig. 93) and details of the accessory genitalia
(fig. 48) of the holotype, and the vulvar lamina of the
allotype (fig. 172).
Male (n=6, including holotype). — Venational sta-
tistics. Fifth (rarely sixth) antenodal thickened; num-
ber of marginal cells behind fore wing paranal cells:
0/0; anx: fore wing 13-18/15-17, hind wing 10-12/9-
12; pnx: fore wing 11-13/15-16, hind wing 11-
14/11-13; cs under pterostigma: fore wing 5-6/6-7,
hind wing 5-7/6-7; anal triangular cells 3-4/3-4.
Hind wing 26-27 mm. Abdomen 30-33 mm.
Female (n = 4, including allotype). — Venational
statistics . Fifth or sixth (rarely fourth) antenodal
thickened; number of marginal cells behind fore wing
paranal cells: 0-1/0-1; anx: fore wing 15-17/15-16,
hind wing 11-12/9-12; pnx: fore wing 11-15/13,
hind wing 10-12/11-13; cs under pterostigma: fore
wing 6-7/6-7, hind wing 6-7/6-7. Hind wing 28-29
mm. Abdomen 31-32 mm.
Diagnosis
Erpetogomphus sabaleticus is diagnosed under E.
tristan.
Remarks
Variation. — Venational details of holotype male:
number of marginal cells behind fore wing paranal
cells: 0/0; anx: fore wing 15/16, hind wing 11/11;
pnx: fore wing 13/13, hind wing 12/11; cs under pte-
rostigma: fore wing 6/6, hind wing 6/6; anal triangu-
lar cells: 3/3. Hind wing 26 mm.
Venational details of allotype female: number of
marginal cells behind fore wing paranal cells: 1/0;
anx: fore wing 16/15, hind wing 12/11; pnx: fore
wing 12/13, hind wing 10/11; cs under pterostigma:
fore wing 6/6, hind wing 6/7. Hind wing 29 mm.
The living colours of the holotype male were re-
corded by Williamson (1918).
Biology. — Williamson (1918) collected the allo-
type female and holotype male along the upper San
Juan near Maraquita in Colombia. The male was tak-
en along the Quebrada Sabaleticus as it rested on a
flat leaf about five feet above the water. Only one oth-
er specimen of this species was seen (but not collect-
ed) during their stay in Colombia. A description of
the type locality is given by Williamson (1918).
Michael May collected one male and three female
larvae of the species at the Quebrada Juan Grande,
along Pipeline Road in the Panama Canal Zone on
28 January 1975 and 23 January 1977. These were
brought back to Gainesville, Florida, where the adults
emerged about three months later.
This species, like E. tristani, is apparently rare.
Other odonatists and I have collected along Pipeline
Road in Panama, but have failed to find the species.
195
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Distribution (fig. 198). — Erpetogomphus sabaleticus
is the most austral of the genus and is currently
known from one locality in northwestern Venezuela,
one locality in northern Colombia, and Panama. It is
apparently the only member of the genus found in
the first two countries. Its northern limit is unknown,
but it is probably parapatric with the more northerly
E. tristani. Collection dates range from 3 February
(Maraquita, Colombia) through 24 June (Panama
Canal Zone).
Material
Type data. — Holotype male: CoLomBia: Dept.
Antioquia: Cristalina, 28 km on railroad above
PueromBernoMml meh la MER de:
Williamson); allotype female: CoroMmB1A: Dept.
Tolima: Maraquita, 3 Feb. 1917 J. H. and E. B.
Williamson). Both specimens in UMMZ.
Other material (68, 49, including holotype 3
and allotype 9). — PANAMA: Canal Zone. Quebrada
Juan Grande, Pipeline Road, 28 Jan. 1975 (M. L.
May), 12 larva, emerged 19 June 1975 (rsca); same
data but 23 Jan. 1977 (Fsca), 16, 29 larvae, male
emerged 16 May 1977, females emerged 20, 26 May
1977; Pipeline Road, 1.7-4.8 mi NW of Gamboa,
2nd bridge, 24 June 1970 (E. S. Morton), 1d (twp);
VENEZUELA: Tachira Dept.: Tachira, 11 April 1920
(J. H. and E. B. Williamson, W. H. Ditzler), 36 (24
in UMMZ, ld in FSCA).
Erpetogomphus tristani Calvert
(figs. 3-thx, 49-hamules, 72-penis, 94-app, 120-ept,
146-vertex, 173,174-vl, 198-distr)
Erpetogomphus tristani Calvert, 1912a: 290 (descr. of d, 2).
— Calvert 1912b: 384 (mentions mating adaptations);
Williamson 1918: 1 (comparison with E. sabaleticus); Ris
1917: 154 (Panama); Ris, 1918: 154 (Panama); Calvert
1920b: 339 (mentions mating adaptations); Williamson
and Williamson 1930: 12 (summary of status); Paulson
1982: 256 (Costa Rica, Panama); Davies and Tobin
1985: 28 (cat.); Tsuda 1986: 87 (cat.); Bridges 1991:
VII.212 (cat.); Tsuda 1991: 95 (cat.); Belle and Quintero
1992: 100 (key, Panama).
Description
Male. — A thorough description of the holotype
male and allotype female is given by Calvert (1912). I
illustrate the caudal appendages of the holotype (fig.
94) and vulvar lamina of the allotype (fig. 173). In
addition to Calvert’s description, I add the following:
A strong ventral tooth at base of cercus (plainly seen
in lateral view, fig. 94), which in most specimens is
partially hidden by lateroposterior margin of tergite
10; tips of epiprocts narrow in posterior view (fig.
120); anterior hamule black, divided at apical 0.25,
lower branch a rounded shoulder; anterior (upper)
196
branch well developed; posterior hamule pale, acutely
triangular, with a prominent apical tooth; penis with
well developed prepuce, which, when viewed lateral-
ly, is hidden by ventrolateral margin of segment 4; la-
teral lobe small, rudimentary, barely visible, and with
spinulose margin; cornua rudimentary, forming an
ental membranous plate with a quadrate margin (fig.
72). Abdomen 31-33 mm.
Venational statistics. — (n = 5, including holotype).
Fifth antenodal thickened; number of marginal cells
behind fore wing paranal cells: 0-1/0; anx: fore wing
13-15/14-15, hind wing 10-12/10-11; pnx: fore wing
11-13/11-13, hind wing 11-13/10-12; cs under pte-
rostigma: fore wing 5-6/5-6, hind wing 5-6/5-6; anal
triangular cells: 3/3. Hind wing 25-27 mm.
Female. — Vulvar lamina (fig. 174, allotype; 173)
broadly connected, relatively unspecialized, and with
a gently U-shaped notch; basal plate poorly devel-
oped; juncture of Y-shaped postlamellar ridge posteri-
or to posterior margin of lamina; well defined arcuate
depression on either side of central stem. Abdomen
31-33.5 mm.
Venational statistics. — (n = 4, including allotype).
Fifth antenodal thickened; number of marginal cells
behind fore wing paranal cells: 0-1/0; anx: fore wing
15-16/15-17, hind wing 11-12/10- 12; pnx: fore
wing 12-14/11-12, hind wing 12-13/12-13; cs under
pterostigma: fore wing 5-7/6-7, hind wing 6-7/5-6.
Hind wing 29 mm.
Diagnosis
Erpetogomphus tristani, E. sabaleticus, and E. cons-
trictor form a compact group which is easily separable
from all other congeners by the unique shape of the
epiprocts of the male (figs. 92-94, 120) and by the
morphology of the vertex of the female (figs. 145-
146). Males of E. tristani differ from the other two
species only in the shape of the cerci. The ventral
margin is distinctly convex at the apical 0.60-0.80 in
E. tristani (fig. 94) and E. sabaleticus (fig. 93), not
smoothly concave as in E. constrictor (fig. 92). All th-
ree species have a prominent dorsal tooth near the
distal end, but the tooth in £. constrictor is broader in
lateral view and more vertical than in £. tristani and
E. sabaleticus.
Males of E. tristani and E. sabaleticus approach one
another closely. Williamson (1918: 4-5) found males
of both species inseparable except by the male caudal
appendages. Williamson sent a pair of E. sabalecticus
to Calvert, who also supported the statement of dif-
ferences between the males of both species. Calvert
was unable to separate the females of these two spe-
cies. I have found the females of these three species
(E. constrictor, E. tristani and E. sabaleticus) to be ex-
tremely similar, more so than their males. The sides
of the longitudinal trough of the vertex of E. constric-
tor are more parallel (fig. 145), than those of the oth-
er two species. Like Calvert and Williamson, I have
not found any differences between females of the
small series of E. tristani and E. sabaleticus.
Remarks
Variation. — Little variation is expressed among the
small series (males, n = 5; females, n = 4, including
holotype and allotype).
Venational details of holotype male: number of
marginal cells behind fore wing paranal cells: 0/0;
anx: fore wing 14/14, hind wing 11/11; pnx: fore
wing 12/12, hind wing 13/12; cs under pterostigma:
fore wing 5/5, hind wing 5/5. Hind wing 27 mm.
Venational details of allotype female: number of
marginal cells behind fore wing paranal cells: 0/0;
anx: fore wing 15/15, hind wing 11/11; pnx: fore
wing 14/12, hind wing 13/12; cs under pterostigma:
fore wing 7/7, hind wing 7/6. Hind wing 29 mm.
Biology. — The species is relatively rare in collec-
tions, despite intensive collecting in Costa Rica in re-
cent years. Collection dates are February, June, and
July. Nothing else is known of its biology.
Distribution. (fig. 198). — Erpetogomphus tristani is
thus far known from only two northwestern provinc-
es of Costa Rica and Panama (Lino, Chiriqui Prov. at
about 1600 m, 8°48’N, 82°26W, 16, 1 ©, Ris
1917). It is replaced to the north by £. constrictor (fig.
198) and to the south by Æ sabaleticus.
Erpetogomphus tristani will probably be found in
southern Nicaragua, but its southernmost distribu-
tion is unknown.
Material
Type data. — Holotype male with following label
data (all handwritten by P. P. Calvert unless otherwi-
se noted): ‘Oricuajo,/ Costa Rica./ July, 1911/ J. F.
Tristan’, ‘Erpetogomphusl tristanil 3/ Calvert/ Tyre’,
‘Needham/ fig’ [in Needham’s hand?], red label:
‘9244 Type [printed] 944/ Erpetogomphusl tristani 6 |
Calvert.’, ‘2nd ham-/ ules/ sheath of/ penis.’ Allotype
female with same label as holotype except for follo-
wing: ‘Labium &/ Maxillae.’, “Erpetogomphusl trista-
ni Q/ Calvert’. Both specimens were originally pin-
ned, but each has been placed in a clear envelope.
Both specimens in ANSP.
Other material (54,49, including holotype d and
allotype ®). — Costa Rica: Guanacaste Prov.: Nicoga,
Feb. 1912 (J. F. Tristan), 18 (ummz); Quebrada
Azul, 2.5 mi W of Tilaran, 24 July 1967 (O. S. Flint,
Jr., and Ortiz B.), 1d, 12 (usnm); Rio Santa Rosa,
3.7 mi E of Las Canas, 25 July 1967 (O. S. Flint, Jr.,
and Ortiz B.), 12 (usnm); Puntarenas Prov.: 8 mi
WNW of Esparta, 26 June 1967 (O. S. Flint, Jr., and
Ortiz B.), 1¢ (usnm); stream 8.1 mi WNW of
GARRISON: Revision of Erpetogomphus
Esparta, 26 June 1967 (O. S. Flint, Jr., and Ortiz B.),
16 (pre); San Jose Prov.: Res. Biol. El Rodeo, 7 km W
of Villa Colon, 9°54’N 84°16°W, 800 m, 10-13 July
1990 (T. W. Donnelly), 19 (twp).
Erpetogomphus ophibolus Calvert
(figs. 4-thx, 29-abd, 50-hamules, 73-penis, 95-app,
121-ept, 147-vertex, 175-vl, 198-distr)
Erpetogomphus ophibolus Calvert, 1905: 163 (Mexico:
Atoyac in Veracruz, H. H. Smith, 24). — Calvert 1909:
489 (distr., Mex.); Muttkowski 1910: 87 (cat.); Calvert
1912a: 294 (comparison with £. tristanı); Ris, 1917: 155
(comparison with Æ. constrictor); Williamson and
Williamson 1930: 489 (summary of status); Kimmins
1969: 296 (type in BMNH); Montgomery 1973: 239 (der-
ivation of name); Paulson 1982: 255 (Mex.); Davies and
Tobin 1985: 27 (cat.); Tsuda 1986: 87 (cat.); Bridges
1991: VII.154 (cat.); Tsuda 1991: 95 (cat.).
Erpetogomphus (Erpetocyclops) ophibolus. — Carle 1992: 148
(key to subgenera, type species of Erpetocyclops subg. n.).
Description
Male. — Labium, labrum, mandibles, anteclypeus
brown; postclypeus entirely brown, or with small la-
teral green triangular spot; frons green except for
brown along frontoclypeal suture, and at base of an-
tefrons; vertex dark brown; area anterior to median
ocellus undifferentiated, postocellar tubercles well de-
veloped, connected medially by a low transverse rid-
ge, slightly tuberculate medially; occiput planar, tra-
pezoidal, hind margin linear or slightly concave
medially, crest covered with long brown hairs; post-
occiput brown, transversely concave; rear of head
brown.
Prothorax brown, anterior lobe green, posterior
lobe green brown; synthorax (fig. 4) with prominent
well defined dark stripes on green background as fol-
lows: middorsal stripe slightly narrowing dorsally, an-
tealar sinus and area along this structure connecting
to humeral stripe, antehumeral stripe of same width
as humeral connecting ventrally and (but not always)
dorsally, and in some specimens broadly so, antehu-
meral not or just touching antealar sinus; second and
third lateral stripes connecting at upper 0.75, isolat-
ing round green spot below subalar carina; posterior
margin of metepimeron, metasternum and venter of
thorax grey brown; venter of profemora grey green,
remainder of femora dark brown, becoming black
distally; tibiae, tarsi, armature black. Wings hyaline,
venation and pterostigma black.
Venational statistics. Fifth (rarely fourth or sixth)
antenodal thickened; number of marginal cells be-
hind fore wing paranal cells: 0/0; anx: fore wing 13-
15/13-16, hind wing 10-12/10-12; pnx: fore wing
10-13/8-12, hind wing 9-12/9-12; cs under pteros-
tigma: fore wing 4-7/5-8, hind wing 5-6/4-7; anal tri-
197
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
angular cells: 3/3. Hind wing 25-27 mm.
Abdomen (fig. 29) with segment 1 brown, green
posterolaterally and with a green middorsal stripe;
segment 2 brown except for green auricles, ventral
margin, and narrow middorsal stripe; annulus dark
brown; segment 3 primarily dark brown, becoming
black distally with green middorsal stripe extending
from annulus of segment 2 and narrowing posterior-
ly to end, a pale grey anterolateral spot becoming nar-
rower along ventral margin and disappearing at 0.25
to 0.50 of segment length; segments 4-6 similar to
segment 3 but with pale middorsal stripe ending at
posterior 0.10 to 0.75 of segment and anterolateral
pale spot ending at transverse carina; segment 7 grey
green on anterior 0.50 to 0.60, except for dark trans-
verse carina, posterior part of segment brown to
black; segments 8-10 red brown, becoming darker
dorsally and ventrally, especially along foliate mar-
gins; posterior medial margin of segment 9 bluntly
pointed or slightly so. Abdomen 30-33 mm.
Cercus (fig. 95) ivory brown, becoming brown dis-
tally, arcuate with a prominent laterally compressed
dorsal tubercle on distal 0.75, extreme base with a
black ventral tooth (mostly hidden by posterior mar-
gin of abdominal segment 10), though not as promi-
nent as in £. constrictor, a ventral carina on distal 0.30
of cercus, tip of cercus with a black tooth; epiprocts
brown, gently curved at distal 0.50, tips slightly di-
verging, each forming a blunt point (fig. 121), a well
defined anterodorsally projecting tooth at middle of
medial margin of epiproct.
Accessory genitalia. Hamules (fig. 50) small; ante-
rior hamule brown, divided at distal 0.30, posterior
branch a well developed shoulder; posterior hamule
roughly triangular, its tip with a spine; penis (fig. 73)
with cornuae slightly bilobate or transverse at base; a
well developed prepuce, lateral lobe a small tooth.
Female. — Head markings similar to male but with
some green on mediolateral areas of labrum, brown at
base of antefrons with a median extension connecting
with frontoclypeal suture, thus dividing frontal green
into two spots. Morphology of vertex and occiput
similar to male with following differences: transverse
postocellar ridge (fig. 147) complete, slightly arcuate
medially, with a slight shallow notch laterally before
resuming at postocellar tubercle; occiput a narrow
planar semicircle, crest covered with long brown
hairs; postocciput tumid, brown; rear of head brown.
Pro- and synthorax as in male, with dark markings
in some specimens less extensive.
Venational statistics (n = 11). Fifth (rarely fourth
or sixth) antenodal thickened in all wings; number of
marginal cells behind fore wing paranal cells: 0-1/0-2;
anx: fore wing 14-16/14-17, hind wing 10-12/10-13;
pnx: fore wing 10-13/9-13, hind wing 10-13/10-14;
cs under pterostigma: fore wing 5-8/5-8, hind wing
198
5-7/4-7. Hind wing 26-29 mm.
Abdomen with segment 1 brown, a green spot pos-
terolaterally and a complete green middorsal stripe;
segment 2 brown with following areas green: wide
middorsal stripe narrowing posteriorly, incomplete
lateral stripe extending from just anterior to auricle to
black posterior annulus, a thin pale margin at venter
of segment; segment 3 similar to segment 2 but
brown becoming darker; middorsal stripe narrowing
posteriorly and forming a mere line on posterior 0.25
of segment; lateral pale stripe ending posteriorly at
transverse carina; segments 4-6 similar to segment 3
but middorsal green becoming increasingly smaller
on successive segments, so that only a small middor-
sal spot occupies basal 0.10-0.50 of segment 6; seg-
ment 7 with anterior 0.50 pale green white, interrupt-
ed by brown transverse carina, posterior 0.50
becoming red brown to almost black; segment 8-10
all red brown to dark brown, darker dorsoposteriorly;
cerci pale grey green. Vulvar lamina (fig. 175) with
plates broadly connected at basal 0.75; relatively un-
specialized, cleft obtusely V- or U-shaped, about as
wide as each lobe; postlamellar ridge with Y- juncture
posterior to plates, base of stem well defined and with
oval depression laterally. Abdomen 30-33 mm.
Diagnosis
Males of E. ophibolus are unique in possessing an
anterodorsal spine on the mediodorsal surface of the
epiproct (fig. 95). The cercus of this species is most si-
milar to that of E. constrictor (fig. 92). Morphology of
the vertex and occiput serve to distinguish female £.
ophibolus from similarly marked species. The median
ocellus is in line with the lateral ocelli (fig. 147), sepa-
rating this species from E. sabaleticus, E. tristant (fig.
146), and E. constrictor (fig. 145). Female E. ophibolus
more closely resemble females of E. schausi, E. agki-
strodon, and E. eutainia. However, E. schausi and E.
agkistrodon have a transverse postocellar ridge which is
medially bilobed (figs. 148-149), while E. eutainia
lacks any postocellar ridge and has postocellar tuber-
cles only. The postocellar ridge in E. ophibolus is arcu-
ate and entire medially (fig. 147). Erpetogomphus op-
hibolus and E. eutainia are often sympatric, and the
females superficially resemble one another. The se-
cond and third thoracic stripes in E. eutainia are al-
ways separate (figs. 7-9), but are connected at their
upper 0.30 in E. ophibolus (fig. 4). Finally, females of
E. eutainia lack the basal stem of the postlamellar rid-
ge (fig. 178) present in E. ophibolus (fig. 175).
Remarks
Variation. — Most specimens of E. ophibolus that I
examined are from southern Veracruz, and they show
little variability in maculation. The single male from
Chiapas has slightly more extensive black thoracic
markings: the pale area between the dark humeral and
antehumeral stripes is a narrow line. The two females
from Belize have the apical dark brown on abdominal
segment 7 laterally extending anteriorly separating
the lateral pale area into one anterodorsal and one an-
teroventral spot. The acuminate condition of the pos-
terior medial margin of abdominal segment 9 varies
intraspecifically.
Biology. — This small green species often occurs
with £. eutainia at small rivulets along the lowland
Gulf of Mexico drainage systems of eastern Mexico.
In 1976 I collected many by flushing them from agri-
cultural stubble bordering trees and shrubs along the
Rio Otapa in central Veracruz. Adults, when flushed,
did not fly far and were consequently easy to collect.
Six other gomphids collected at the same site under
the same circumstances included Phyllocycla brevi-
phylla Belle, P. volsella (Calvert), Phyllogomphoides
suasus (Selys), P. duodentatus Donnelly, Progomphus
clendoni Calvert, Erpetogomphus eutainia, and E.
bothrops.
Raúl López (in litt.), from information given to
him by Enrique González, writes that E. ophibolus is
the most common gomphid in Veracruz, found year-
round in the vicinity of the Los Tuxtlas Biological
Station near Catemaco. Altitudinal range is from sea
level to 615 m (Chiapas). Collection dates range from
8 June (Belize) to 13 September (Veracruz).
Distribution (fig. 198). — This species is thus far
known from southern Veracruz (southeast of 19°N,
97°W), central Chiapas, Guatemala, and Belize.
Tineke Boomsma (in litt.) collected a pair of this spe-
cies in the Mountain Pine Ridge area of Belize (Cayo
Distr.: Privassion Creek, 24 July 1993).
Material
Type data. Holotype male: [Mexico] Vera Cruz,
Atoyac, May (H. H. S[mith])/ Erpetogomphus ophi-
bolus Calv. Tyre d. P. P. Calvert det. 1905. B. C. A.
Neur., p. 164. Original of pl. 7, figs. 30-32, 46
(Kimmins 1969). In BMNH.
Other material (436, 122). — Mexico: Veracruz:
Rio Otapa, 8 km S of La Tinaja, 90 m, 13 Aug. 1976,
(R. W. and J. A. Garrison), 188, 32, (RwG); 20 Aug.
1976, 123, 12 (Rwe); Playa Escondida, 20-24 June
1981, (E. Gonzalez), 12, (UNAM); Arroyo cerca de
Playa Escondida, 17-24 July 1981, (E. Gonzalez), 19,
(UNAM); arroyo nr. Playa Escondida, about 30 km NE
of Catemaco, 18°34-36°N, 95°04-09°W, 23 July-6
Aug., 9-13 Aug. 1982, (R. W. Garrison), 5d, (RWG);
Los Tuxtlas, arroyo despues de Laguna Escondida, 25
May 1980, (E. González), 16, (swp); 11 June 1980,
(R. Novelo), 1d, (unam); Los Tuxtlas, stream nr.
Jicacal Beach, 10 July 1979, (Gerado Jimenez), 19,
(cc); Colonia Apachital, 16 km S, 10 km E of Tierra
Blanca, 10 Sept. 1965, (T. W. Donnelly), 2d, 19,
GARRISON: Revision of Erpetogomphus
(TWD, CC, UNAM); Rio Hondo, on rd. to “Colonia la
Apachital’ 50 ft [15 m], 6 mi E of Hwy from Tierra
Blanca to Cd. Aleman, 25-26 Aug. 1957, (G. H.
Beatty III), 1d, (esca); 3 km N of Santiago Tuxtla, 13
Sept. 1965, (T. W. Donnelly), 12, (cc); Chiapas.
stream 20.1 mi N Ocozocoautla, 2000 ft [615 m], 25
Aug. 1967, (D. R. and M. L. Paulson), 1d, (rsca);
Berrize: Toledo Distr: Blue Creek Village, EARTH-
WATCH Belize Expedition, 1981, 8 June 1981, (D. H.
Messersmith, W.E. Steiner, et al), 19, (usNM); Cayo
Distr. Mountain Pine Ridge, Rio Frio at Augustine,
16°58’N, 88°59’W, 500 m, 22-25 July 1983, (T. W.
Donnelly), 19, (rwp); GUATEMALA: El Progreso Dept:
6.5 km N of Est. de la Virgén, 29 Aug. 1965, (T. W.
Donnelly), 14, 22, (rwp).
Erpetogomphus agkistrodon sp. n.
(figs. 5-thx, 30-abd, 148-vertex, 176-vl, 198-distr)
Type material. — Holotype 2, Mexico: Veracruz
State, Parque Javier Clavijero, Jalapa, 23 Aug. 1982,
1300 m (R. López) in UNAM.
Description
Male. — Unknown.
Holotype female. — Labium pale grey, becoming
dark grey medially; labrum, clypeus, base of mandible
dark brown washed with dark olive green along me-
dian area of labrum, ante-clypeus, and above lateral
lobes; pale green spot along lateral margin of labium;
broad green stripe above frontoclypeal suture contin-
uing to near base of vertex; antennae, vertex, occiput
dark brown; transverse postocellar ridge (fig. 148)
complete, emarginate medially; rear of occiput broad-
ly semicircular; rear of head dark brown.
Prothorax entirely brown except for green anterior
lobe; synthorax (fig. 5) with dark brown thoracic
stripe well developed on green background; triangular
middorsal stripe interrupting inverted green ‘7’ at
base, upper end of middorsal stripe connecting with
antehumeral and humeral stripes, these two united
leaving only pale spot below antealar sinus, and small
green streak below; second lateral stripe connecting
with humeral and third lateral stripes above and be-
low, this stripe also connecting with third lateral be-
low antealar sinus, thus isolating a green spot; meta-
spiracle black; posterior margin of metepisternum
with a dark brown stripe; metasternum dark brown-
grey. Femora red-brown, becoming black distally; re-
mainder of legs and armature black.
Wings hyaline, venation and pterostigma black.
Venational details. Sixth antenodal thickened in all
wings; number of marginal cells behind fore wing par-
anal cells: 3/2; anx: fore wing 15/17, hind wing 12/11;
pnx: fore wing 14/16, hind wing 15/15; cs under pt:
fore wing 6/5, hind wing 5/6. Hind wing 31 mm.
199
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Abdomen (fig. 30) predominantly black with ves-
tiges of a narrow green middorsal stripe on segments
2-6; anterior 0.25 of segment 7 green, becoming
white laterally; broad brown midlateral stripe on seg-
ment 2 becoming darker on segment 3 and more ex-
tensive on succeeding segments, thus encircling small
white anterolateral spots; auricle on segment 2 green;
segments 8-10 entirely black, cercus white. Vulvar
lamina as shown in fig. 176. Abdomen 31 mm.
Diagnosis
Erpetogomphus agkistrodon belongs to the E. trista-
ni group because of the simple condition of the vulvar
lamina and the complete postocellar ridge. This spe-
cies is distinguished from females of all other species
by the emarginate condition of the postocellar ridge.
That of E. ophibolus is entire. The female of E. agki-
strodon is similar to the only known female of £.
schausi, and is diagnosed under the latter species.
Remarks
I suspect the male of E. agkistrodon will have a pe-
nis similar to that of E. ophibolus or E. schausi; it
should have a prepuce and the lateral lobes should be
small, semi-circular and with or without spinules.
The thickened sixth antenodal in this species is an
unusual condition: the fifth antenodal is usually
thickened throughout the genus. However, this may
be an anomaly of the holotype.
No conspicuous depressions or pits are present an-
terior to the median ocellus. This area, instead, forms
a shallow transverse V. Based on female morphology,
I suspect the male epiprocts will be relatively thick,
gently curved at posterior 0.5, forming a 90° angle,
tip broadly spatulate.
Biology. — The female was collected along a shaded
creek in company with Cordulegaster diadema godma-
ni McLachlan. Erpetogomphus boa were also taken at
the locale, though they were flushed from low vegeta-
tion about 50 m from the stream.
Distribution (fig. 198). — Known only from the
type locality.
Erpetogomphus schausi Calvert
(figs. 6-thx, 31-abd, 51-hamules, 74-penis, 96-app,
142-face, 149-vertex, 177-vl, 198-distr)
Erpetogomphus schausi Calvert, 1919: 33 (desc. holotype d).
— Calvert 1920a: 113 (note on type); Williamson and
Williamson 1930: 13 (summary of status); Paulson 1982:
255 (Guat.); Davies and Tobin 1985: 28 (cat.); Tsuda
1986: 87 (cat); Bridges 1991: VII.187 (cat.); Tsuda
1991: 95 (cat.).
Description
Male. — A thorough description of the holotype
male accompanies the original description. I illustrate
200
the thorax (fig. 6), abdomen (fig. 31), accessory geni-
talia (fig. 51), penis (fig. 74), head (fig. 142), and cau-
dal appendages (fig. 96) of the holotype. The speci-
men has been skewered with a bristle. The cercus of
the holotype (fig. 96) is strongly curved at distal 0.30,
the ventral margin has a remnant of an inferior carina
at its extreme base, but is well developed again on the
distal 0.50.
Venational statistics (based on holotype d and one
3 from Costa Rica): number of marginal cells behind
fore wing paranal cells: 3/2-3; anx: fore wing 16-
18/16-20, hind wing 12-14/12-13; pnx: fore wing
15/14-15, hind wing 13-14/13-15; cs under pteros-
tigma: fore wing 5-6/5-6, hind wing 6/5-6; anal trian-
gular cells: 4/4. Hind wing 29-30 mm. Abdomen 32-
33 mm.
Female. — Labium grey; labrum brown with indica-
tion of green mediolaterally, ante- and postclypeus
brown with possible green medially on postclypeus;
frons green with brown along frontoclypeal suture
and base of antefrons, especially medially, with off-
shoot almost touching frontoclypeal suture, thus al-
most separating green; base of antefrons with a slight
longitudinal raised area with a slight concavity on
each side; antennae, vertex brown; transverse post-
ocellar ridge (fig. 149) complete, emarginate medial-
ly; occiput brown, semicircular crest covered with
long brown hairs; postocciput brown, convex, rear of
head brown.
Prothorax entirely brown, paler on anterior and
posterior lobes; synthorax as in holotype male (fig. 6);
femora grey brown, tibiae and tarsi darker, armature
black.
Wings hyaline, venation and pterostigma black.
Venational statistics (n = 1). Fifth antenodal thick-
ened in all wings; number of marginal cells behind
fore wing paranal cells: 2/0; anx: fore wing 16/16,
hind wing 12/11; pnx: fore wing 15/15, hind wing
14/14; cs under pterostigma: fore wing 6/5, hind
wing 6/6. Hind wing 31 mm.
Abdomen compressed laterally with markings
mostly unrecognizable, but probably similar to £. ag-
kistrodon (fig. 30). Vulvar lamina simple (fig. 177),
similar to that of E. agkistrodon. Abdomen about 32
mm.
Diagnosis
Erpetogomphus schausi is unique in possessing a
combination of characters of the penis (with well de-
veloped prepuce) and caudal appendages (with gently
decumbent cercus (fig. 96)). Although the morpholo-
gy of its penis allies this species with E. sabaleticus, E.
tristani, and E. constrictor, the penes of the other three
species lack well developed cornuae and a spinulate
lateral lobe, typical of £. schausi. The only known fe-
male differs from all other known species except E.
agkistrodon in possessing a complete transverse posto-
cellar ridge which is notched medially. The only
morphological difference that I have noted between
these two species is the broader, more semicircular oc-
ciput (figs. 149) of E. schausi, compared with the nar-
rower, more broadly semicircular condition found in
the holotype of E. agkistrodon (fig. 148). The sixth
antenodal crossvein is thickened in all four wings of
E. agkistrodon, while the fifth is thickened in this fe-
male of E. schaust; but, as stated under E. agkistrodon,
I suspect the condition in the holotype is atypical.
Remarks
Variation. — The male sex of this species is known
from only two specimens, the holotype male descri-
bed in detail by Calvert (1919), and another male fr-
om Costa Rica collected by J. Belle and loaned to me
for inclusion in this paper. The Costa Rican male is
slightly smaller (HW = 29 mm) than the holotype
(HW = 30 mm) and differs as follows (condition for
holotype in parentheses): small green triangle just la-
teral to medial area of anteclypeus (all dark brown);
round green spot below antealar sinus of mesepister-
num smaller (bigger); anterior pale abdominal spots
extending well beyond transverse carina on segments
3-7 (these spots smaller and confined to anterior 0.25
of segments 4-6, to 0.5 of segment 7); sides of abdo-
minal segments 8-10 light brown (dark brown); later-
al lobe of penis more angulate (more rounded); se-
cond thickened antenodal in fore wing 5-5 (6-7) and
in hind wing 5-6 (6-6).
The holotype appears to be a fully mature male:
traces of pruinosity are present on the venter of its
thorax. The Costa Rican male seems to be less ma-
ture: it lacks pruinosity, and the dark brown areas of
the holotype are pale in this specimen. The green ar-
eas of the thorax of the Costa Rican male are slightly
more restricted than for the holotype.
The teneral female described above seems to repre-
sent the female of this species. The combination of
characters allying it with the £. tristani group and the
occurrence of a male from Puntarenas Province, lead
me to suspect that I have associated this specimen
correctly.
Before the discovery of the two £. schausi from
Costa Rica, I considered the possibility that Æ. agkis-
trodon might be the female of E. schausi: morpholog-
ical considerations in light of the characters for other
members of the £. tristani group (e.g., vulvar lamina,
shape of occiput) supported this view. However, the
geographic distance (about 1700 km), coupled with
the local occurrence of both forms, tend to discount
this. It is possible that the morphological differences I
detect in the occiput of female E. agkistrodon and E.
schausi, though major, may be due to geographic vari-
ation, which would render E. agkistrodon a junior
GARRISON: Revision of Erpetogomphus
synonym of E. schausi.
Biology. — Nothing is recorded of the circumstanc-
es of capture of the holotype. Jean Belle records the
following for the Costa Rican male: “This gomphid
was secured on the grass-covered bank of an almost
impassable mountain rivulet where the insect alighted
after a clash with a Hetaerina specimen. The colors
when alive are as follows: compound eyes dark
brown; head and thorax dark brown with light green;
middorsal pale spots of abdominal segments 3 to 6
and pale spot of abdominal 7 yellow.’
Altitudinal data range from 1220 m to 1500 m; the
female was collected 8-9 April, the Costa Rican male
on 30 August.
Material
Type data. — Holotype male: GUATEMALA: Dept.
Purulh4, Baja Vera Paz, 1220 m, forest stream (W.
Schaus, J. Barns), 7 July [no date], in ANsp.
Other material (2d, 1 2, including holotype à). —
Costa Rica: Puntarenas Prov: Monte Verde, 1500
m, rivulet, 30 Aug. 1986 (J. Belle), 16 (RNHL); Rio
Bellavista, ca. 1.5 km NW of Las Alturas (8.951 N,
82.846 W), elev. 1400 m, 8-9 April 1987
(Holzenthal, Hamilton, Heyn), 12 (usNM).
Distribution (fig. 198). — Known only from the
type locality in Guatemala and Puntarenas Province,
Costa Rica. It probably occurs in intervening Central
American republics.
Erpetogomphus eutainia Calvert
(figs. 7, 8, 9-thx, 32, 33-abd, 52-hamules, 75-penis,
97-app, 122, 123, 124, 125-ept, 143, 144- face,
150-vertex, 178-vl, 199-distr)
Herpetogomphus menetriesii. — Selys, 1878: 429 (24 sep.)
(Guatemala. (Un mâle [sic, female] incomplet. Musée de
Paris, une femelle. — Coll. Selys.’)).
Erpetogomphus eutainia Calvert, 1905: 162 (1d, Rio
Papagayo in Guerrero, Mexico). — Calvert 1919: 35
(comparison with E. schaust); Kimmins 1969: 293 (type
in BMNH); Paulson 1982: 255 (U.S., Mex., Guat.);
Davies and Tobin 1985: 27 (cat.); Tsuda, 1986: 87 (cat.);
Bridges 1991: VII.72 (cat.); Tsuda 1991: 95 (cat.).
Erpetogomphus diadophis Calvert, 1905: 167 (26, Texas). —
Muttkowski 1910: 87 (cat.); Needham and Heywood
1929: 79 (descr.); Needham and Westfall 1955: 147
(descr.); Borror 1963: 104 (common name);
Montgomery 1968: 133 (distr.); Kimmins 1969: 293
(type in BMNH); Paulson, 1982: 266 (synonymy of E. di-
adophis and E. eutainia); Bridges 1991: VII.60 (cat.).
Erpetogomphus ? diadophis. — Calvert 1919: 36 (Guat., pos-
sible conspecificity with E. diadophis)
Herpetogomphus diadophis. — Byers 1928: 5 (larva un-
known).
Erpetogomphus (Calogomphus) eutainia. — Carle 1992 (key to
subgenera, type species of Calogomphus subg. n.).
201
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Description
Male. — Labium grey to dark grey, labrum all green
with a small medial spot on anterior margin (fig. 144)
to entirely brown with large central green spot (fig.
143), base of mandibles green, tips becoming black,
anteclypeus all green to largely brown; postclypeus gr-
een with brown medially along frontoclypeal suture
and descending laterally along lateral lobes, thereby
isolating medial green spot; in more southerly speci-
mens, brown expanding so that green confined to la-
teral margins; frons green with brown along fronto-
clypeal suture and at extreme base of antefrons; vertex
brown with medial transverse area at postocellar tu-
bercles green, to entirely brown; occiput brown,
transverse triangular pits anterolateral to medial ocel-
lus, postocellar tubercles low, incomplete medially;
occiput trapezoidal, largely planar except for medial
swelling; crest costate, linear to slightly concave, rim-
med with long brown hairs; postocciput green me-
dially to all brown, concave; rear of head brown.
Prothorax brown along middle lobe, remainder
green or becoming entirely brown except for pale an-
terior lobe; synthorax (figs. 7-9) with prominent,
well-defined dark brown stripes on green background
as follows: middorsal stripe expanding toward collar,
divided by pale middorsal thoracic carina, not touch-
ing collar, medial 0.5 or entire antealar crest, linear
antehumeral, not touching upper margin; humeral,
second and third lateral stripes, medial area of mesin-
fraepisternum; more southerly specimens with these
stripes becoming thicker (figs. 8-9) so that antehu-
meral and humeral connect at lower and upper ends;
a vestigial stripe along posterior margin of metepime-
ron; venter of thorax and metasternum pale grey
green to grey. Coxae, trochanters pale green, venter of
profemora green, remainder of profemora black; met-
afemora dark brown to black except for pale area ven-
trally; basal 0.40 to 0.60 of metafemora pale green to
dark grey, becoming black apically; tibiae, tarsi and
armature black.
Wings hyaline, venation and pterostigma black;
more northerly specimens with pale, narrow line
along costa disappearing proximal to pterostigma.
Venational statistics. Fifth, rarely fourth, antenodal
thickened in all wings; number of marginal cells be-
hind fore wing paranal cells: 0-1/0-1; anx: fore wing
11-15/11-15, hind wing 8-11/8-10; pnx: fore wing 8-
11/8-12, hind wing 8-11/8-11; cs under pterostigma:
fore wing 4-6/4-6, hind wing 4-6/4-6; anal triangular
cells: 3-4/3-4. Hind wing length 23-25 mm.
Abdomen (fig. 32) with segment 1 green and a pale
brown dorsolateral stripe on each side; segment 2
brown except for green auricles, ventrolateral margin,
and middorsal stripe; segment 3 with lateral brown
stripe, interrupted medially just posterior to trans-
verse carina, and with an irregular pale spot within
202
anterior brown spot, posterior brown more promi-
nent posteriorly and meeting dorsally on posterior
0.10 of segment; more southerly specimens with lat-
eral brown not interrupted medially and with pale an-
terolateral spot reduced or almost absent; incomplete
middorsal stripe blue green, pale areas becoming
white laterally; segments 4-6 similar to segment 3 but
with dark brown more prominent posteriorly and
with anterolateral white spot confluent with ventral
margin; segment 7 pale blue green on anterior 0.50-
0.60, except orange brown transverse carina, posteri-
or part of segment dark red brown; segments 8-10 red
brown, becoming darker dorsally and ventrally, espe-
cially along foliate margins; posterior medial margin
of segment 9 smoothly carinate, or with a moderately
blunt point. Abdomen 29-32 mm.
Cercus (fig. 97) ivory, becoming brown distally,
gently arcuate at distal 0.30, with a small black tooth,
a ventral costate carina on basal 0.35-0.40 of cercus,
often ending in a small, isolated, pebble-like tooth; in
other specimens, the posterior part of this carina may
be pebble- like; epiprocts (figs. 122-123) brown,
moderately curved along posterior 0.50, tip of epi-
proct in lateral view thick, bluntly pointed, with lat-
eral costate margin; tips in posterior view (figs. 124-
125) planar, roundly divergent.
Accessory genitalia. Hamules (fig. 52) small, ante-
rior hamule brown, distal 0.25 divided, the superior
branch meeting the inferior branch, thus enclosing an
oval space; posterior hamule small, digit-shaped, no
apical hook; penis (fig. 75) with long pointed cornu-
ae; dorsal membranous hood prominent, its ends par-
allel; lateral lobes large, frill-like, their margins with
spinules; dorsal area of membranous third segment
with a pair of mediolateral blunt chitinized tubercles.
Female. — Head as in male with following differ-
ences: brown at base of antefrons projecting anterior-
ly, partially isolating dorsal green; vertex morphology
similar to male, postoccipital tubercles small, isolated
medially, occiput trapezoidal as in male, planar with
faintly convex arcuate crest; postocciput brown, pla-
nar.
Thorax as in male.
Venational statistics. Fifth, rarely sixth, antenodal
thickened in all wings; number of marginal cells be-
hind fore wing paranal cells: 0-2/0-3; anx: fore wing
13-16/13-16, hind wing 9-11/8-11; pnx: fore wing 8-
12/8-11, hind wing 8-12/8-13; cs under pterostigma:
fore wing 4-7/4-7, hind wing 4-7/4-7. Hind wing 25-
28 mm.
Abdomen (fig. 33) with segment 1 green with a
brown dorsolateral stripe; segment 2 similar to seg-
ment 1, but with wash of brown along ventral margin
of tergite; segment 3 with dorsolateral stripe darker,
expanding posteriorly and connecting above at poste-
rior 0.10 to 0.20 or separated by thin hairline of
green, ventral margin with brown interrupted by
white below brown dorsolateral stripe; in more north-
erly specimens, connecting above with middorsal
green stripe; posterior part of brown connecting with
dorsolateral stripe posteriorly, transverse carina dark
brown; segments 4-6 similar to segment 3 but brown
darker, more definite, connecting posterodorsally on
0.25-0.50 on progressive segments, lateral white con-
fined to spots, one at anteroventral margin, another
ventromedially, these two spots sometimes connected
by narrow line of white ventrally; more northerly
specimens with medioventral white connecting with
middorsal green stripe; segment 7 with anterior 0.50
white except for darkened transverse carina, posterior
0.50 dark brown; segments 8-10 dark brown; cerci
white, their tips black. Vulvar lamina (fig. 178) with
plates small, almost digit-shaped; U-shaped cleft
about as wide as or wider than each plate; postlamel-
lar ridge with a costate semicircular ridge. Abdomen
29-33 mm.
Diagnosis
The characteristically shaped posterior hamules
(fig. 52) and penile structures (short prepuce, wide
spinulate lateral lobe [fig. 75]) separate males of this
species from all others except E. leptophis.
Erpetogomphus eutainia possesses a straight or pebble-
like ventral carina on the basal 0.25 of the cercus (fig.
97); no such carina is present in E. leptophis (figs. 98).
Females of E. eutainia and E. leptophis are unique
in possessing a semicircular postlamellar ridge; in all
other congeners, this structure is a Y-shaped ridge.
The posteriorly pointed occipital protuberance in E.
leptophis (fig. 151) is distinct from the relatively un-
modified condition present in £. eutainia (fig. 150).
Remarks
Variation. — Venational details of holotype male of
E. eutainia. No marginal cells behind fore wing pa-
ranal cells; anx: fore wing 14/14, hind wing 10/10;
pnx: fore wing 10/9, hind wing 10/10; cs under pte-
rostigma: fore wing 5/5, hind wing 5/6; number of
anal triangular cells: 4/4. Hind wing 24.5 mm.
Venational details of holotype female of £. diado-
phis. number of marginal cells behind fore wing para-
nal cells: 0/1; anx: fore wing 13/14, hind wing 10/10;
pnx: fore wing 10/9, hind wing 10/11; cs under pte-
rostigma: fore wing 6/7, hind wing 6/5. Hind wing
28 mm.
This small species has a widespread distribution
and, consequently, exhibits some degree of clinal vari-
ation north to south. Specimens from southeastern
Texas and northern Mexico are the palest: the dark
transverse markings of the face are narrower, and
green is often present medially on the vertex and oc-
ciput. The labrum is mostly green (fig. 144), the dark
GARRISON: Revision of Erpetogomphus
thoracic stripes are narrow, and none is present on the
posterior margin of the metepimeron (fig. 7). Medial
bands of white encircle abdominal segments 3-6 in
both sexes. The dark somatic markings are darker,
broader, and more definite in specimens from south-
ern Veracruz. Vestiges of a stripe on the posterior
margin of the metepimeron can be seen (fig. 8), and
the pale white medial abdominal bands are obscured,
entirely or almost entirely interrupted by the dorso-
lateral brown stripe. Specimens from Costa Rica are
the most melanic (figs. 9, 143); no green is present on
the vertex or occiput, and the middorsal green stripe
of abdominal segments 3-6 is always separate from
the medioventral white.
The ventral carina of the cercus is also variable.
Some specimens have a continuous costate carina
with a small, isolated, pebble-like tooth distally; but
others may have several antepenultimate pebble-like
teeth in addition to the longer ultimate one. The tips
of the epiproct may also be variable. In most speci-
mens, these structures are posteriorly flattened (fig.
122) with the outer margin slightly to strongly con-
cave (tig. 0125) iphemaleiitromy El Progresoy
Guatemala, has the epiprocts similar to the condition
described above, but they are narrower (fig. 125).
The clinal differences described above probably led
Calvert (1905) to interpret the female of E. diadophis
as a separate species from E. eutainia. Calvert (1919)
later received another female from Guatemala, which
he postulated may be the same as the paratype of E.
diadophis, he made no mention of E. eutainia.
The large series of males and females examined
from various parts of its range convince me that
Paulson (1982) was correct in synonymizing the two
names. The status of the two females (not male and
female) described by Selys (1878) as Herpetogomphus
menetriesii is discussed under the status of E. mene-
triesit. 1 also examined a female labelled
‘Cyanogomphus | mexicanusl n. sp. / coll. R. Martin /
2’, from Honduras, in the MNHP.
Biology
I collected this small, colourful species in southeas-
tern Texas over several years. They were flushed from
stubble bordering agricultural fields next to the
Gonzales River. They were sympatric with the larger
E. designatus, and were about as common. However,
the former species behaved more like a damselfly, for
they never flew far, rested on tips of stubble or barbs of
barbed-wire fences, and were always easy to take with
a net. I collected one male which was the prey of a rob-
ber fly (Asilidae); another, I rescued from an orb-we-
aver spider web. I was able to photograph a pair in co-
pula in the late afternoon.
In southern Veracruz at the Rio Otapa, E. eutainia
had similar habits, as described under £. ophibolus.
203
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
The female from El Salvador (Rio Palio) has the fol-
lowing by V. Hellebuyck: ‘on bushes along a fast run-
ning river, clear water, rocky bottom & shady banks’.
Altitudinal ranges are from near sea level (20 m,
Medina Bank, Belize) to 615 m (Oaxaca). Collection
dates range from 19 May (Costa Rica) through Oc-
tober (Veracruz, La Gloria; Guerrero, Rio Papagaio).
Distribution (fig. 199). — This species occurs
southeast of the Edwards Plateau in southern Texas,
south along the lowlands and foothills of the Sierra
Madre Oriental in northern and central Mexico. The
farthest western locality is El Sabino in Michoacan
state. Specimens have been collected sporadically in
Belize, Guatemala, Honduras, El Salvador, and
northern Costa Rica.
Material
Type data. — Holotype male of E. eutainia [labels
all printed unless otherwise stated]: ‘R. Papagayo, /
Guerrero, 1200 ft. / Oct. H. H. Smith.’, ‘Brit. Mus. /
1911-339., ‘ERPETOGOMPHUS/ eutainia Calvert
[written]/ P. P. Calvert, det. 1905 [written]/ B. C. A.
Neur., p. 162 [written]/ Original of Pl. 7, f. 24-27, 39
[all written]’, small round label with red border:
‘Holo-/ type’, in BMNH. Holotype female (not male,
as erroneously stated by Kimmins, 1969) of E. diad-
ophis. [labels all printed unless otherwise stated]:
‘Texas [written by an unknown hand]’, pale violet la-
bel: ‘McLachlan Coll./ B. M. 1938-674.’, ‘ERPETOG-
OMPHUS 9 [written]/ diadophis Calv. Tyre [written]/
P. P. Calvert, det. 1905 [written]/ B. C. A. Neur., p.
167 [written]/ original of Pl. VII, ff. 35, 47 [all writ-
ten]’, in BMNH.
Material examined (74d, 429, including holotypes of £.
eutainia and E. diadophis). — U. S. A.: Texas: Caldwell Co.:
San Marcos R. at Luling, (esca, cc); Gonzales Co.:
Guadalupe River, 4 mi S of Gonzales, (cc, Rw@); near
Otting, Palmetto State Park, (rsca); Palmetto State Park, 6
mi S of Luling, (RwG); Mexico: Michoacan: El Sabino, 20 mi
SSE of Uruapan (UMMZ, Rw); Oaxaca: La Escondida, route
190, km 727, 2000’, (usnm); San Luis Potost: Tamazunchale,
(usnm); Tamaulipas. 3 mi S of Ciudad Victoria, (USNM);
Veracruz. Cardel, La Gloria, (UsNM); pond 2.7 mi S of La
Tinaja, 300 ft., (DRP); Rio La Palma, 25 km N of Catemaco,
(rwG); Rio Otapa, 8 km S of La Tinaja, 90 m, (rw);
Sontecomapan, Coscoapan, (UNAM); BELIZE: Toledo Dist.:
Medina Bank, 20 m, (TB); GUATEMALA: El Progreso Dept.:
San Agustin Ac., (esca); Zacapa Dept.: Zacapa, (ESCA); un-
known locality and date (‘Rodig.’ [illegible]) [labelled as
Herpetogomphus menetriesii), (IRSN); unknown locality, date,
and collector [labelled ‘E. menetriesi? Selys], (MNHP);
Honpuras: Unknown locality, date, and collector [labelled
‘Cyanogomphus?/ mexicanusl n. sp./ (Coll. R. Martin/ 97],
(MNHP); EL SALVADOR: La Libertad Dept.: Rio Palio, San
Juan Opico, 400 m, (va); Costa Rica: Alajuela Prov.:
Quebrada Mina 0.2 mi S Hwy. 11 on Hwy, (rsca);
Guanacaste Prov: Hda. Taboga, 100 ft., (pre); Rio Santa
Rosa, 3.5 mi N of Canas, 300 ft (pre).
204
Erpetogomphus leptophis sp. n.
(figs. 10-thx, 34, 35-abd, 53-hamules, 76-penis, 98-
app, 151-vertex, 179-vl, 200-distr)
Type material. — Holotype male. BeLIZE: Toledo
District, I. Z. E. field station, Blue Creek, 25 June
1983 (M. L. May) (rsca). Allotype female. BELIZE:
Toledo District, Blue Creek Village (Earthwatch)
Belize Expedition 1981, 8 June 1981 (D. H.
Messersmith, W. H. Steiner, et al.) (USNM).
Description
Holotype male. — Labium light grey, becoming
dark grey medially; labrum green with dark brown
margin and medial line almost separating green into
two spots; base of mandibles grey-green, anteclypeus
green, postclypeus brown except for small green spots
above lateral lobes; ventral margin of frons brown,
joining brown of postclypeus, thus forming a large
frontoclypeal stripe; remainder of frons green; a large
transverse furrow at base of antefrons, this area with a
pair of deeper pits, each anterolateral to median ocel-
lus; postocellar ridge complete, but with medial part
lower and at level of lateral ocelli; extreme base of
frons, entire vertex, and occiput dark brown, occiput
tumid medially, hind margin mostly straight but pe-
aked medially so that hind margin seems slightly an-
gularly convex; rear of head dark brown.
Prothorax entirely brown except for following
green areas: anterior lobe, and two small midlateral
spots (almost touching) on median lobe; synthorax
(fig. 10) with dark brown thoracic stripes well devel-
oped on green background; dark middorsal stripe tri-
angular, its wide base touching collar but not con-
necting with antehumeral; antehumeral and humeral
connected along upper 0.25, thus isolating small
green spot; second lateral as shown in fig. 10, a poste-
rior branch connecting with third lateral along upper
0.30, isolating a green spot; posterior margin of met-
epimeron brown, all thoracic stripes connected by
brown along antealar sinus; metasternum grey-
brown. Venter of profemora grey-green, remainder of
legs dark brown; tibiae, tarsi, armature black.
Wings hyaline, venation and pterostigma black.
Venational details. Fifth antenodal thickened in all
wings; no supplementary marginal cells behind fore
wing paranal cells; anx: fore wing 14/13, hind wing
10/10; pnx: fore wing 10/11, hind wing 10/10; cs
under pterostigma: fore wing 6/5, hind wing 5/5;
anal triangular cells: 4. Hind wing 29 mm.
Abdomen as in fig. 34. Segment 1 brown, green
laterally and with a green middorsal spot; segment 2
brown except for green auricles, ventral margin, and
narrow middorsal spot with a cordate expansion in
the middle; segment 3 black with green middorsal
stripe ending at basal 0.30 of segment and grey ante-
rolateral spot; segment 4 black with anterior 0.2
white laterally, becoming green dorsally; segments 5-
G like segment 4; segment 7 with anterior 0.5 white
laterally, becoming green dorsally, posterior 0.5 of
segment dark red-brown; segments 8-10 red-brown,
posterior medial margins of segments 8-9 slightly
pointed. Abdomen 31 mm.
Cercus (fig. 98) light green, simple, with no ventral
carinae, slightly arcuate, tip black; epiprocts brown,
gently curved at distal 0.5, tips viewed posteriorly
with parallel sides, forming bluntly rounded tips.
Accessory genitalia. Hamules (fig. 53) small; ante-
rior hamule brown, distal 0.25 divided, the superior
branch meeting the inferior branch, thus enclosing an
oval space; posterior hamule small, finger-like, no ap-
ical hook; penis (fig. 76) with long, pointed cornuae,
dorsal membranous hood long, its ends crossing (fig.
76); lateral lobes large, frill-like, their margins with
spinules.
Allotype female. — Head as in male except on la-
brum, brown medial line separating green into two
spots; base of antefrons as in male (fig. 151) with a
raised bilobed ridge between lateral ocelli; posterior
margin of occiput with a prominent, posteriorly di-
rected, pointed protuberance, its base tumid so that
entire raised area assumes shape of a plumbob.
Thorax as in male.
Wings hyaline, similar to male.
Venational details. No supplementary marginal
cells behind fore wing paranal cells; anx: fore wing
14/16, hind wing 9/11; pnx: fore wing 11/13, hind
wing 10/11; cs under pterostigma: fore wing 5/4,
hind wing 5/6. Hind wing 25 mm.
Abdomen (fig. 35) with segment 1 brown, becom-
ing dark green-brown laterally; segment 2 brown with
a narrow green middorsal stripe and green lateral
stripe encircling auricle and extending entire length
of segment; segment 3 like segment 2, but brown be-
coming darker and green lateral stripe abbreviated,
forming an elongate spot along basal 0.5 of segment;
segments 4-6 all black with white basal band, extend-
ing ventroposteriorly along ventral margin of tergites;
segment 7 with anterior 0.30 white, remainder dark
red-brown; segments 8-10 dark red-brown, cercus
white. Vulvar lamina (fig. 179) with plates connected
anteriorly, plates wider than in £. eutainia; postlamel-
lar ridge semicircular as in £. eutainia. Abdomen 30
mm.
Diagnosis
The structure of the hamules and penis ally £. lep-
tophis with E. eutainia, but the superior appendages
easily distinguish the two species (figs. 97-98).
Erpetogomphus eutainia has a pebble-like ventral cari-
na on the basal 0.25 of the cercus; but no such struc-
ture is present in £. leptophis. The posteriorly directed
GARRISON: Revision of Erpetogomphus
pointed occipital protuberance easily identifies the fe-
male of E. leptophis. The female shares with E. eutai-
nia the semicircular postlamellar ridge, but E. leptop-
his possesses a bilobed ridge between the lateral ocelli.
This surface is planar in £. eutainia.
Remarks
Although the types were taken by different collec-
tors two years apart, the female characters indicate it
is the female of E. leptophis.
Biology. — Michael May collected the male along a
creek in the forest.
Distribution (fig. 200). — Known only from the
type locality.
Erpetogomphus elaphe sp. n.
(figs. 11-thx, 54-hamules, 77-penis, 99-app, 180-vl,
200-distr)
Erpetogomphus elaps. — Calvert 1907: 398 (Costa Rica, fig-
ures anterior hamules).
Erpetogomphus sp. n. near elaps. — Dunkle 1988: 46
(Honduras).
Description
Holotype male. — Labrum pale grey green; antecly-
peus, labrum, base of mandibles pale green, tips of
mandibles dark brown; postclypeus and frons pale gr-
een, a slight wash of brown on lateral margins of fron-
toclypeal suture, antefrons pale green, slightly darker
at base, vertex dark brown; a deep transverse trough
in front of median ocellus; pedicel, scape, and flagel-
lum brown; occiput trapezoidal, pale green, mostly
planar, slightly tumid medially, crest slightly promi-
nent, slightly emarginate medially and covered with
long brown hairs; postocciput green, transverse when
viewed dorsoposteriorly; rear of head yellow brown.
Prothorax green, becoming brown on middle lobe
laterally and anteromedially. Synthorax (fig. 11) ap-
ple green with following dark areas: poorly defined
middorsal stripe not extending to collar and inter-
rupted medially by middorsal thoracic carina and an-
tealar crest, well defined antehumeral stripe narrow-
ing dorsally but touching antealar crest and extending
ventrally to mesinfraepisternum, a small posterior off-
shoot from antehumeral stripe at upper 0.10 connect-
ing with vestigial first lateral stripe which is abbreviat-
ed to upper 0.50; venter of synthorax green, coxae
and trochanters grey green, femora pale green becom-
ing dark brown on distal extensor surfaces, these dark
markings occupying distal 0.80 of profemora, about
distal 0.60 of mesofemora, and distal 0.30 of metafe-
mora; tibiae and tarsi black; armature black.
Wings hyaline, anterior margin of costa yellow to
pterostigma, thereafter black; yellow at base of costa
darkening to brown at costal triangle, pterostigma
205
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
brown, veins bordering it black.
Venational details. Fifth antenodal thickened in all
wings; number of marginal cells behind fore wing
paranal cells: 1/1; anx: fore wing 13/13, hind wing
10/10; pnx: fore wing 9/9, hind wing 9/10; cs under
pt: fore wing 6/5, hind wing 4/6; anal triangular cells:
4/3. Hind wing 26 mm.
Abdomen with segment 1 yellow green with a wash
of brown dorsolaterally; segment 2 yellow green dor-
sally and ventrally, with a brown midlateral stripe ex-
panding ventrally behind auricle, anterior 0.5 of this
stripe poorly defined, annulus dark brown; segment 3
light yellow green dorsally interrupted laterally by a
longitudinal brown stripe well marked at transverse
carina, expanding again at posterior 0.25 of segment
and connecting dorsally at black annulus; inferior
margin of tergite probably ivory (post mortem preser-
vation has partially obscured this area); segments 4-6
similar to segment 3, with anterior 0.10 of dark mid-
lateral interrupted and with gradual dorsolateral ex-
pansion of stripe so that inverted dorsal pale green
wedge-shaped middorsal stripe disappears at distal
0.40 of segment 6; segment 7 pale green brown be-
coming tawny on posterior 0.10 of segment; seg-
ments 8-10 red brown becoming darker dorsolateral-
ly on segment 8; foliate expansion and denticles
black. Abdomen 33 mm.
Cercus (fig. 99) pale ocher becoming dark brown
at tip; linear, gradually narrowing distally, tips round-
ly pointed, a weakly defined ventral carina on distal
0.50 of cercus; epiprocts brown, gently curved at 90°
angle; tips spatulate when viewed posteriorly.
Accessory genitalia (fig. 54). Anterior hamule green
basally, becoming dark brown distally, branched at
distal 0.50; superior (upper) branch greatly enlarged,
as long as stem of hamule with resulting interval
between branches small, semicircular; posterior ha-
mule (fig. 54) almost digit-shaped, wider at base, with
a poorly developed anterior shoulder, tip broadly
rounded, without an apical tooth; penis with a prom-
inent knife-like serrated lateral lobe; membranous
hood well developed but lobes not overlapping, cor-
nua pointed but with median shoulder as in fig. 77.
Allotype female. — Similar to male in markings and
colouration. Head with transverse trough anterior to
median ocellus as in male, but occipital region differ-
ing as follows: occiput small, erect, barely visible in
dorsal view, convex medially; postocciput with me-
dian tumid area. Thoracic, leg, and abdominal pat-
terns as in male, but dark areas more restricted; mid-
dorsal and antehumeral stripes faint, first lateral stripe
vestigial, confined to upper 0.20 of thorax.
Venational details. Fifth antenodal thickened in all
wings; number of marginal cells behind fore wing
paranal cells: 1/2; anx: fore wing 14/15, hind wing
10/10; pnx: fore wing 10/11, hind wing 11/12; cs
206
under pterostigma: fore wing 6/5, hind wing 5/6.
Hind wing 29 mm.
Vulvar lamina (fig. 180) with basal plate promi-
nent, lobes narrow with medial margins distinctly
concave, their bases not touching; distal part of lobe
with a ventral flap bent under; its costate rim visible
externally and disappearing medially near base of lobe
(fig. 180); with resultant cleft forming a wide, U-
shaped interval; Y-shaped juncture of postlamellar
ridge not extending beyond posterior margin of plate,
stem with diagonally ovaloid depression on each side.
Abdomen 34 mm.
Diagnosis
Erpetogomphus elaphe males differ from males of
the more northerly £. elaps by only two morphologi-
cal characters (contrasting characters for E. elaps in
parentheses): 1) The superior branch of the anterior
hamule is robust and is as high as the basal 0.5 of the
hamule so that the resulting gap between upper and
lower branches is small (fig. 54), (superior branch less
robust, thinner; resultant gap between hamular bran-
ches wider [figs. 55-56]), and 2) The tip of the poste-
rior hamule is rounded, (tip of posterior hamule with
a cephalad directed tooth). The posterior hamule of
E. elaphe appears to be more digit-shaped and is remi-
niscent of those of E. eutainia and E. leptophis. The
same structure appears more triangular in £. elaps.
However, this last character is subtle and is best de-
tected when specimens of both species are in hand: I
do not consider it a good diagnostic character. I have
found no significant differences in the penis or caudal
appendages between the closely related and allopatric
E. elaphe and E. elaps. I have been unable to find any
differences between females of the two species.
Differences between £. elaphe and other similar
species parallel those of £. elaps and are discussed
under that species.
Remarks
Variation. — Erpetogomphus elaphe does not show as
great variation in thoracic markings as does E. elaps.
The holotype male is the most boldly marked of the
series. All paratype males (n = 15), two of which are
teneral, have an antehumeral stripe, although it is ob-
scure in one male from Costa Rica. The humeral stri-
pe in all but one of the paratypes is restricted to the
upper 0.20 of the humeral suture, and the middorsal
stripe is vestigial or absent.
Female paratypes of £. elaphe are similar in macu-
lation to the allotype, although one female from
Costa Rica has a poorly defined antehumeral stripe.
The small humeral stripe is reduced to the upper 0.20
and barely visible.
Venational statistics for paratype males (n = 18);
number of marginal cells behind fore wing paranal
cells: 0-2/0-3; anx: fore wing 12-15/12-15, hind wing
9-11/9-11; pnx: fore wing 8-11/9-11, hind wing 9-
11/9-11; cs under pterostigma: fore wing 3-6/4-7,
hind wing 4-6/4-7; anal triangular cells: 3-5. Hind
wing 25-28 mm. Abdomen 30-34 mm.
Venational statistics for paratype females (n = 6);
number of marginal cells behind fore wing paranal
cells: 0-4/0-3; anx: fore wing 11-17/14-16, hind wing
10-12/10-12; pnx: fore wing 9-12/10-12; hind wing
9-12/9-13; cs under pterostigma: fore wing 4-6/4-5,
hind wing 5-6/4-6. Hind wing 27-30 mm. Abdomen
33-34 mm.
Biology. — Little is known of the biology of this
species, though its habits probably mirror those of £.
elaps. The female from Cartago Province was collect-
ed in a ‘marshy area by grassy hillside’. Altitudinal
gradients for £. elaphe range from about 900 m (nr.
Camotán, Guatemala) to about 1700 m (SE of
Cartago, Costa Rica). Collection dates range from 2
June (Agua Caliente, Guatemala) to 5 August (San
Jose, Costa Rica).
Distribution (fig. 200). — Erpetogomphus elaphe is
known only from Guatemala, Honduras, and Costa
Rica, but it probably exists in El Salvador and
Nicaragua. Its distribution is allopatric (or possibly
parapatric) with the form of £. elaps with thin anteri-
or hamules.
Material
Type data. — Holotype male: (GUATEMALA,
Guatemala Dept. El Fiscal, 6 June 1909 (E. B.
Williamson). Allotype female: same data, but 4 June
1909. Both in ummz.
Paratypes examined (186, 72): GUATEMALA:
Chiquimula Dept.: Alda dos Quebradas nr. Camotän,
900 m (T. W. Donnelly) , 12 (twp); Guatemala
DEDEN Caliente, 2 junel909) (ESB:
Williamson), 19; El Fiscal, 4 June 1909 (E. B.
Williamson) ee MS Mjune m0) (ENB:
Williamson), 14; 6 June 1909, 25 (all ummz); Finca
El Rosario, 36.5 km SE of Guatemala City, 15 June
1975 (J. E. Hafernik, Jr.), 26 (RwG); HONDURAS:
Francisco Morazan Dept.: 30 km ESE Tegucigalpa, 29
June 1985 (Pinto and Ranch), 19 (swp); 24 July
1983 (L. Cordoba), 14 (swp); COSTA RICA: [no local-
ity data], 1920 (Paul Serre), 38 (MNHP); Cartago
Prov., 5 mi SE Cartago, 5500 ft, 14 June 1963 (F. G.
Thompson), 19 (pre); San Jose Prov.: (all collected
by H. Schmidt), San José, 18 June [no year stated],
16 (H. Kahl Coll’n, Acc. 12676); 24 June, 1d; 4
July, 1d; 5 July, 2d; 17 July, 1d, 19; 18 July, 19;
21 July, 1d; 5 August, 1d (rsca).
Erpetogomphus elaps Selys
(figs. 12, 13-thx, 55, 56-hamules, 78-penis, 100-
app, 152-vertex, 181-vl, 200-distr)
GARRISON: Revision of Erpetogomphus
Erpetogomphus elaps Selys, 1858: 330 (70 sep.) (4 descr. ‘Le
Mexique, d’aprés un exemplaire unique du Museum de
Paris, rapporté par M. Sal[l]e.’). — Selys 1859: 538 (12
sep.) (descr. 3); Selys 1869: 175 (12 sep.) (descr. d, 2);
Selys 1873b: 519 (75 sep.) (list); Calvert 1905: 163 (lo-
calities in Mex.); Calvert 1907: 399 (comparison with E.
boa); Calvert 1909: 481 (distr. in Mex.); Muttkowski
1910: 87 (cat.); Ris 1917: 153 (mentions Calvert’s
[1907] comments on comparison with Æ bod);
Williamson and Williamson 1930: 13 (summary of stat-
us); Montgomery 1973: 239 (derivation of name);
Paulson 1982: 255 (Mex., Guat., Costa Rica); Davies and
Tobin 1985: 27 (cat.); Tsuda 1986: 87 (cat.); Bridges
1991: VII.68 (cat.); Tsuda 1991: 95 (cat.).
Gomphus elaps. — Hagen, 1861: 100 (descr. 3)
Herpetogomphus elaps. — Hagen 1875a: 42 (cat.); Selys 1879:
64 (note on classification); Kirby 1890: 60 (cat.);
Needham 1897: 182 (name mentioned); Calvert 1899:
386 (descr. d, comparison with £. viperinus auct.), 415
(gizzard); Higgins 1901: 128 (mentions Calvert, 1899).
Description
Male. — Labrum grey white with wash of brown
around margin of median lobe; anteclypeus, labrum,
base of mandibles pale grey green, tips of mandibles
dark brown; postclypeus and frons light green, some
specimens with brown at base; vertex dark brown; a
deep transverse trough in front of median ocellus, a
small tubercle behind each lateral ocellus; pedicel,
scape and flagellum brown; occiput trapezoidal, light
green, mostly planar; crest slightly prominent,
straight, slightly notched medially, and covered with
long brown hairs; postocciput green, transverse when
viewed dorsoposteriorly; rear of head yellow brown,
paler laterally.
Prothorax and median lobe red brown becoming
pale apple green medially, especially on anterior and
posterior lobes. Synthorax ranging from entirely apple
green (fig. 13) with slight hint of dark antehumeral
stripe to having following brown areas (fig. 12): poor-
ly defined middorsal stripe divided by pale middorsal
thoracic carina, well-defined antehumeral stripe just
touching antealar crest dorsally and not touching me-
sinfraepisternum ventrally; a vestigial first lateral
stripe confined to upper 0.30-0.40 of suture and
sometimes with a small anterior offshoot touching an-
tehumeral; venter of synthorax, coxae, trochanters,
grey green; femora pale green, becoming dark brown
on distal extensor surfaces; these dark markings occu-
pying distal 0.80 of profemora, distal 0.50 of mesofe-
mora, and distal 0.20 to 0.40 of metafemora; tibiae
and tarsi dark brown to black; armature black.
Wings hyaline, anterior margin of costa yellow to
pterostigma, thereafter black; yellow at base of costa
darkening to brown costal triangle; pterostigma dark
brown, veins bordering it black.
Venational statistics. Fifth antenodal (occasionally
fourth or sixth) thickened in all wings; number of
marginal cells behind fore wing paranal cells: 0-2/0-3;
207
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
anx: fore wing 11-15/12-15, hind wing 9-11/8-11;
pnx: fore wing 8-11/8-11, hind wing 8-11/8-11; cs
under pt: fore wing 4-6/4-7, hind wing 4-6/4-7; anal
triangular cells: 3-5. Hind wing 24-29 mm.
Abdomen with segment 1 yellow green with a wash
of brown dorsolaterally; segment 2 yellow green dor-
sally with a large midlateral brown spot dorsoposteri-
orly to auricle, or, in well- marked specimens, a dis-
tinct dark brown midlateral stripe expanding
ventrally behind auricle, annulus dark brown; seg-
ment 3 broadly light yellow green dorsally interrupt-
ed laterally by a longitudinal brown stripe well-
marked at transverse carina and flaring again at
posterior 0.25 of segment to annulus; in palely
marked specimens, brown midlateral stripe interrupt-
ed behind transverse carina; in well-marked speci-
mens, midlateral stripe broad and meeting dorsally on
posterior 0.10 of segment; inferior margin of tergite
ivory, in heavily marked specimens limited to a trian-
gular spot an anterior 0.10 of segment; segments 4-6
similar to segment 3, but some specimens with dark
midlateral stripe more prominent and not interrupted
behind transverse carina; in heavily marked speci-
mens, midlateral stripe almost black and covering en-
tire segment, except for basal pale annulus at basal
0.10 of segment and dorsal inverted pale green
wedge- shaped middorsal stripe gradually narrowing
posteriorly and disappearing at distal 0.20 of segment
4; these dorsal pale areas lacking on segment 5 and 6;
segment 7 pale green, becoming tawny on posterior
0.10 of segment, or with posterior 0.20 dark brown,
becoming black dorsally, transverse carina brown;
segments 8-10 generally red brown becoming darker
dorsally; in darkly marked specimens becoming al-
most black dorsally; foliate expansion and denticles
black. Abdomen 29-38 mm.
Cercus (fig. 100) pale ocher to light brown, linear,
gradually narrowing distally, the tips roundly point-
ed, a weakly defined ventral carina on distal 0.50 of
cercus; epiprocts brown, gently curved at a 90° angle,
tips when viewed posteriorly slightly spatulate, broad-
ly rounded, or obliquely truncate.
Accessory genitalia (fig. 55). Anterior hamule dark
brown to black, branched at distal 0.30, superior
branch larger than inferior branch; superior branch
moderately thickened in specimens from northern
Mexico south to Chiapas, Mexico; specimens south
of Chiapas (fig. 56) with superior arm of hamule
more slender (see remarks); posterior hamule (figs.
55-56) pale, roughly triangular, with an anterior
shoulder, tip with a tooth pointed cephalad; penis
with prominent, knife-like serrated lateral lobe; mem-
branous hood well developed but lobes not overlap-
ping, cornua pointed but with median shoulder as
shown in fig. 78.
Female. — Similar to male in markings and colour-
208
ation; head with transverse trough anterior to median
ocellus as in male, but occipital region differing as fol-
lows: occiput small, planar, encompassed posteriorly
by arcuate (convex) crest (fig. 152); postocciput with
median tumid area. Thoracic, leg, and abdominal
patterns as in male, but dark areas more restricted, es-
pecially on femora and abdomen; cerci pale.
Abdomen 30-36 mm.
Venational statistics. Fifth (rarely sixth) antenodal
thickened in all wings; number of marginal cells be-
hind fore wing paranal cells: 0-3/0-3; anx: fore wing
12-16/13-16, hind wing 9-12/9-12; pnx: fore wing 8-
11/9-12, hind wing 8-12/9-12; cs under pt: fore wing
5-6/5-7, hind wing 5-7/5-7. Hind wing 26-30 mm.
Vulvar lamina (fig. 181) with basal plate promi-
nent, lobes narrow with medial margin distinctly
concave, their bases not touching, with resultant cleft
forming a wide, U-shaped interval; apical 0.30 of
plate with mesal margin folded ventrally, beneath
main plate, the costate rim sometimes visible along
lateral margin of plate (fig. 181); Y-shaped juncture
of postlamellar ridge not extending beyond posterior
margin of plate, stem with diagonally ovaloid depres-
sion on each side.
Diagnosis
Erpetogomphus elaps and E. elaphe are closely rela-
ted species which are diagnosed under £. elaphe. In
males, the linear cercus of £. elaps (fig. 100) easily dis-
tinguishes it from the decumbent cercus (fig. 102) of
E. bothrops. Body colouration is similar in E. elaps and
E. bothrops, but all E. bothrops have well-marked ante-
humeral and humeral stripes (fig. 15). Many speci-
mens of E. elaps have these markings poorly develo-
ped or nearly absent (fig. 13).
With the exception of £. elaphe, whose females ap-
pear indistinguishable, females of E. elaps are most
similar to E. bothrops. The differences in the vulvar
lamina as discussed by Calvert (1899) are an easy way
to distinguish them. Secondly, the occiput of E. elaps
is small and convexly arcuate (fig. 152); in £. both-
rops, this structure is wider (fig. 154) and not nearly as
arcuate.
Remarks
Variation. — The description is based on 49 males
and 21 females. The holotype is in fair condition, alt-
hough most of dorsum of the prothorax and part of
the mesepisternum have been eaten away by dermest-
ids. It has a completely pale thorax and the black mid-
lateral stripes on abdominal segments 3-6 are inter-
rupted only on the basal 0.10 of each segment. The
hamules (fig. 55) and cerci (fig. 100) show that this
specimen is the apple-green species ranging from
northern Mexico south through Guerrero and into
southern Veracruz.
Venational details of holotype male: number of
marginal cells behind fore wing paranal cells: 0/0;
anx: fore wing 12/12, hind wing 9/10; pnx: fore wing
9/8, hind wing 8/9; cs under pt: fore wing 4/4, hind
wing 5/5; anal triangular cells: 3/4. Hind wing 24
mm. Abdomen 31 mm.
I consider E. elaps to be the most variable of any
species in the genus. Calvert (1907) figured three
forms of the anterior hamules of this species when he
examined a male from San Jose, Costa Rica (= E.
elaphe). His fig. 31 (Guerrero, Rincon) and fig. 32
(Morelos, Cuernavaca) correspond to the holotype of
E. elaps (fig. 55), which is characterized by possessing
a moderately thickened anterior branch whose tip al-
most overlaps the posterior branch when viewed lat-
erally. Calvert’s fig. 30 (Guerrero, Rincon) is typical
of specimens I have seen from Chiapas, Mexico, and
northern Guatemala (fig. 56). The anterior branch
forms a slender hook which rarely overlaps the poste-
rior branch when viewed posteriorly. Figures 33 and
34 of Calvert (1907) represent £. elaphe and are dis-
cussed under that species. Calvert (1907) stated that
he was unable to find any other characters which var-
ied correlatively, and I have found no other characters
to separate E. elaps and the narrow hamular form. I
have not found any intermediate conditions in the
shape of the anterior hamule. Their distributions ap-
pear to be largely allopatric, although Calvert (1907)
figures both forms from Rincon in Guerrero.
Interestingly, the distribution of the form with the
thin anterior hamular branch lies between typical £.
elapsand E. elaphe. Because I have been unable to find
any other diagnostic characters, I am inclined to treat
the Rincon male and others from Chiapas and north-
ern Guatemala as a variant of £. elaps. The divergent
morphology of southernmost E. elaps may represent
character displacement between £. elaps and E. elaphe.
Both males and females show remarkable range in
size and maculation patterns throughout its range.
The smallest specimens examined are from Morelos
(HW: males 23 mm, females 26 mm), the largest
from the Rio Metlac region of Veracruz (HW: males
29 mm, females 31 mm).
Specimens collected at 1220 m in Sinaloa State are
melanic, characterized by pale areas of abdominal seg-
ments 4-6 reduced to basal 0.10 laterally; pale areas
on dorsum of segments 3-6 reduced laterally, this pale
area a thin hairline on segment 5, a basal ring on seg-
ment 6.
I find no correlation of presence or absence of the
dark thoracic stripe to locality. A series of specimens
from Palapita, Nayarit, ranges from an entirely pale
thorax to possessing antehumeral and humeral
stripes. All specimens from Veracruz that I have ex-
amined lack dark thoracic stripes, but all southern
variants with the thin anterior hamular branch from
GARRISON: Revision of Erpetogomphus
Chiapas and northern Guatemala possess them.
Biology. — This small species has habits similar to
those described for £. bothrops. The following field
notes accompany specimens collected by E. B.
Williamson: Nov. 6 (Tepic): “2 Gomphine — from
brushy vegetation between road and river at rapids
above mill’; Nov. 7: ‘Gomphine d — Have seen these
only in bushes along bank of Rio de Tepic — usually
not over foot above ground — resting or flying. One 9
was ovipositing in rather still water below the rapids’;
Nov. 14 (Jalisco: San Diego Rancho): ‘Very hard to
catch in net. Sits tight on rock and lets it [net] pass
over and then flies. Easily approached and I finally
began catching them by dropping the net over their
rock & letting them rise into the bag.’; and Nov. 16
(Jalisco: San Diego Rancho): ‘[d, 9] caught in grass
in banana patch early in AM, sluggish’.
López (in litt.) records it as one of the most com-
mon members of the genus in Mexico, having a wide
tolerance for diverse stream habitats. I have seen no
records of this species taken near sea level, unlike £.
bothrops. Altitudinal records range from 670 m
(Nayarit, Palapita) to about 1500 m (Cuernavaca),
dates of capture from 9 June (Veracruz, Teocelo) to
23 November (Jalisco, San Diego Rancho).
Distribution (fig. 200). — Both this species and £.
bothrops appear to be the most widespread members
of this genus in Mexico. Erpetogomphus elaps occurs in
mesic areas on either side of the Sierra Madre
Oriental and Occidental. It has been collected as far
north as Nuevo León in the east, and southern
Sinaloa in the west, southward through Jalisco,
Morelos, Guerrero, and Veracruz.
Material
Type data. — Holotype male with following data:
small round white label with handwritten ‘2/44’; rec-
tangular manila label with handwritten (in R.
Martins hand) ‘Ophiog. elaps/ d De Selys/
Mexique? a renvoyer.’; large green label with black
border with handwritten ‘E. elaps, Selys/ Mexique’;
small white label with printed “MUSEUM PARIS’; nar-
row red label with printed ‘HOLOTYPE'; white printed
label with “E. elaps HOLOTYPE/ Det. J. Belle, 1977/
No. 2, 1844. Insects [sic] de/ [reverse side] differents
Ordres envoyes/ de Mexique par/ Mr. Ghresbreght !
and in lower right hand corner ‘ne pas/ Sal[l]e! In
MNHP. [he caudal appendages and accessory genitalia
are shown in figs. 55, 78, 100.
Other material (496, 212). — Mexico: Chiapas.
Pacific slope, 800-1000 m), (UsNM); stream 20.1 mi
NE Tapanatepec, Oaxaca, 2700 ft, (rsca); San
Geronimo, Volcan Tacana, 450 m (ummz); Distrito
Soconusco, Finca Juarez, (UMMZ); Guerrero:
Chilpancingo, 4600’, (esca, BMNH); Tepetlapa, 3000
ft, (BMNH); Jalisco: San Diego Rancho nr. Cocula,
209
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
(ummz); 15 mi on road to Tequila, (UMMZ);
Michoacan: El Sabino, 20 mi SSE of Uruapan,
(ummz); Morelos: Alpuyeca, S of Cuernavaca on Rt.
95, (rsca); Cerro de Higuerón, Jojutla, (UNAM);
Cuautla, (ummz); Cuernavaca, ca. 1200-1500 m,
(UMMZ, USNM, RWG); 5 mi S of Cuernavaca, (UMMZ);
16 mi S of Cuernavaca, (UMMZ); Itzamatitlan, 5 km
SW of Yautepec, (RwG); Nayarit: Acaponeta,
(BMNH); Jumatán, (UNAM); Tepic, (UMMZ, USNM);
Jalisco: Palapita, 670 m (UNAM); Nuevo León:
Municipio de Monterrey, Cerro El Mirador, (rsca);
Sinaloa: stream 27.9 mi NE of Concordia, 4000 ft.,
(DRP, ESCA, CC, RWG); Veracruz: Barranca de
Cayoapa, Teocelo, (UNAM); Jalapa, (USNM); 4.9 km N
of Coscomatepec by Mex. Hwy 156 (rwe); Rio
Metlac, 2 km WNW of Fortin, 900 m, (Rwo);
GUATEMALA: Suchitepequez Dept: Finca Moca
Grande, Rio Bravo, (rsca).
Erpetogomphus liopeltis sp. n.
(figs. 14-thx, 36-abd, 57-hamules, 79-penis, 101-
app, 117-cercus, 126-ept, 140, 141-occiput, 153-
vertex, 182-vl, 201-distr, 227-wings)
Erpetogomphus elaps. — Novelo & Pena, 1991: 130 (misiden-
tification).
Type material. — Holotype male: Mexico: Nuevo
Leon State, Municipio de Monterrey, La Estanzuela,
2 July 1987 (M. J. Westfall, Jr, H. Quiréz, A.
Contreras). In rsca. Allotype female: Nuevo Leon
State, Monterrey, El Diente, 5 mi SE, 2 July 1960
(W.B. Cutts). In uMMz.
Paratypes examined (15d, 3 9, including holotype
3 and allotype 2). - Mexico: Hidalgo: Pemuxtitla,
Rio Zacuala, 1000 m, 22 April 1984 (R. Novelo), 1d
(RWG); Michoacán: 12 km W Apatzingán, 369 m
(1200 ft.), 4th Hoogstraal Mexican Biological
Expedition, 12 Aug. 1941 (Harry Hoogstraal), 19
(ummz); Nuevo Leon: Chipingue, 12 July 1960 (W.B.
Cutts), 12 (rwG); Municipio de Monterrey, Cerro
El Mirador, 9 Aug. 1984 (A. Contreras), 2d (tori);
same data as holotype, 3d (rsca); Municipio de
Santiago, El Cercado, Arroyo Dolores, 17 Aug. 1984
(A. Contreras), 26 (tori); Municipio de Santiago,
Rancho Los Pinos, 3 July 1987 (M. J. Westfall, Jr.,
Jr, G. Luna, A. Contreras), 56 (rsca, RwG); San Luis
Potosi. La Conchita (Camino a Xilitla), 7 May 1950
(no collector), 16 (cc).
Description
Holotype male. — Entire head pale green, labrum
dull grey green, base of mandible, anteclypeus dull gr-
een, tip of mandible black; basal 0.25 and all of vertex
brown, a transverse trough at anterior margin of ver-
tex at juncture of base with antefrons, this trough
210
with two slightly deeper pits, one each anterolateral to
median ocellus; occiput with slightly medial tumid
area, occipital crest distinctly emarginate medially
(figs. 140-141), dark brown, rimmed with long dark
brown hairs; postocciput green, becoming black late-
rally, concave medially; rear of head brown, beco-
ming yellow brown laterally; antennae dark brown.
Prothorax brown, anterior and posterior lobes and
small dorsolateral area of median lobe pale green.
Thorax (fig. 14) entirely green (probably blue green
in life) with following dark brown: obscure middorsal
stripe ending before collar, becoming darkest near
middorsal thoracic carina, lateral margins of this
stripe gradually fading to green on mesepisternum;
posterolateral rim of antealar crest; lateral 0.40 of
mesepisternum except for dorsal emargination, with
no indication of pale green separating coalesced hu-
meral and antehumeral stripes; this stripe narrowing
slightly ventrally but connecting with lateral margin
of collar and anterior 0.50 of mesinfraepisternum;
rim below subalar carina at obsolete second lateral su-
ture; narrow partial third lateral stripe extending ven-
trally to 0.50 of suture before merging with green.
Metasternum pale grey brown, tawny anteriorly.
Coxae, trochanters grey; base of femora brown, be-
coming black distally; tibiae and armature black.
Wings hyaline, anterior margin of costa dark
brown at base, becoming yellow distally to proximal
level of pterostigma, remainder of venation and pte-
rostigma black.
Venational details. Fifth antenodal thickened in all
wings; no supplementary marginal cells behind fore
wing paranal cells; anx: fore wing 14/15, hind wing
10/10; pnx: fore wing 8/9, hind wing 9/9; cs under
pterostigma: fore wing 7/6, hind wing 6/5; anal trian-
gular cells: 4/3. Hind wing 31 mm.
Abdomen (fig. 36) with segment 1 green dorsally,
paler laterally with a midlateral stripe of brown ex-
panding anteriorly at base and darkening posteriorly
at articulation point; segment 2 similar to segment 1
but with a ventral stem of brown behind auricle from
dark midlateral stripe, midlateral brown becoming
wider and darker along posterior 0.50 of segment, an-
nulus black; segment 3 with pale green middorsal
stripe acuminate and disappearing at posterior 0.10
of segment, black midlateral stripe expanding slightly
at transverse carina and again widening at posterior
0.10 of segment, this area with a ventral offshoot ter-
minating anteriorly at 0.20 of segment; segments 4-6
similar to segment 3, but with middorsal pale green
stripe shorter and narrower on each succeeding seg-
ment, thus black midlateral stripe connecting dorsal-
ly at posterior 0.40 of segment 6, these midlateral
stripes interrupted at anterior 0.10 of each segment
with small longitudinal wedge-shaped white area
along ventral margin of tergite behind transverse cari-
na; segment 7 with basal 0.60 pale grey green, except
for irregular diffuse black spot at transverse carina, re-
mainder of segment red brown, becoming black dor-
sally; segments 8-10 red brown, becoming darker
dorsally; carinae and lateral foliations black, posterior
medial margin of segment 9 pointed.
Cercus (fig. 101) pale ocher, decumbent at posteri-
or 0.30, largely parallel in lateral view except for dis-
tal 0.20, tip with a shiny black tooth; black inferior
carina on distal 0.25; epiproct brown, about 0.45 as
long as cercus, U-shaped in lateral view, tip of epi-
proct when viewed posteriorly (fig. 126) broadly
truncate. Abdomen 40 mm.
Accessory genitalia (fig. 57). Anterior hamule dark
brown, branched at distal 0.30, superior branch larg-
er than inferior branch; posterior hamule pale, rough-
ly triangular with a well developed anterior shoulder,
distal 0.75 curved posteromedially with a well devel-
oped black tooth twisted laterally from broad axis of
hamule; penis with cornuae acuminate as shown in
fig. 79, lateral lobe prominent, knife-like, serrated
posteriorly.
Allotype female. — Similar to male with following
differences: labrum slightly darker (possibly due to
post mortem effects); transverse trough at juncture of
antefrons and vertex well defined with slightly larger
and deeper pit anterolateral to median ocellus; occi-
put strongly reduced to a small, gently convex, large-
ly perpendicular medial arch; postocciput tumid me-
dially, dark middorsal stripe vestigial, with only a
wash of brown on either side of lower arms of antealar
sinus, a narrow poorly defined incomplete green
stripe separating dark brown antehumeral and hu-
meral stripes up to dorsal 0.75. Abdominal segment 3
with acuminate black spot posterolateral to black dor-
solateral stripe projecting anteriorly to black trans-
verse carina; dorsolateral black stripes on segments 4-
7 more extensive; pale middorsal stripe reduced to
basal 0.75 (segments 4-5) to basal 0.50 (segment 7),
and anteriorly touching annulus on each segment;
segments 8-9 black dorsally, red brown laterally.
Abdomen 38 mm.
Vulvar lamina (fig. 182) small, cleft larger than
each plate, each plate connected basally, each with a
raised shelf occupying anterior 0.5, postlamellar ridge
with Y-shaped juncture posterior to hind margin of
lamina, central stem with a well defined circular de-
pression on either side.
Venational details. Fifth antenodal thickened in
left fore wing; the sixth in remaining wings; Number
of marginal cells behind fore wing paranal cells: 2/3;
anx: fore wing 16/17, hind wing 13/11; pnx: fore
wing 10/12, hind wing 12/11; cs under pterostigma:
fore wing 6/7, hind wing 6/7. Hind wing 34 mm.
GARRISON: Revision of Erpetogomphus
Diagnosis
This remarkable species approaches £. viperinus in
size, colour, and maculation, but is most closely rela-
ted to E. bothrops in morphology. Males of E. liopeltis
and £. bothrops are similar in possessing epiprocts with
broadly truncate tips (figs. 126-128); but those of E.
bothrops are usually, but not always, bidentate (fig.
128). In lateral view, the epiprocts of £. liopeltis are
more strongly curved and extend about 0.5 or less the
length of the cercus; the same structures in £. bothrops
extend 0.75 or more the length of the cercus (fig. 102).
The apical tooth of the posterior hamule of E. Lopeltis
is longer and not as tightly recurved as in E. bothrops
(figs. 57-58). Abdominal segments 1-3 in E. hopeltis
(fig. 36) are more heavily marked than in £. bothrops
(fig. 37); and the dark thoracic markings are also mo-
re extensive (fig. 14) than in E. bothrops (fig. 15).
In size and overall colouration, £. /iopeltis seems in-
distinguishable from £. viperinus but differences in
the tips of the epiproct (figs. 126, 129) and structure
of the area anterior to the vertex in both sexes (figs.
136 [as for E. bothrops|, 137) easily distinguish the
two species. Subtle differences between these two spe-
cies exist in the morphology of the distal 0.5 of the
cercus. In E. liopeltis, these structures are thicker and
more cylindrical when viewed dorsally (fig. 101) and
mediodorsally (fig. 117). The cercus of E. viperinus is
more laterally compressed (fig. 119). The tip of the
cercus in E. liopeltis terminates in a sharp black tooth;
this tooth is lacking in E. viperinus.
Females of E. liopeltis and E. bothrops can be separ-
ated by overall size (E. liopeltis hind wing 32-34 mm;
E. bothrops hind wing 28-31 mm) and thoracic mac-
ulation and colouration, as stated for males.
Structural differences of the occiput and postocciput
easily distinguish these species. The occiput of £. lio-
peltis is rudimentary, with only a gently rounded ver-
tical medial crest. The medially tumid postocciput is
easily visible in dorsal view (fig. 153). In E. bothrops,
the occiput is narrow, but the anterior and posterior
margins are roughly parallel except for the angulate
lateral margins (fig. 154). In E. bothrops, the crest is
vertical and the exposed postocciput is easily visible;
but its posterior margin is planar or only slightly tu-
mid. Receptacles for the tips of the male cerci differ
slightly between the females. The transverse trough
located anterior to the median ocellus narrows lateral-
ly in E. bothrops (fig. 136); but the lateral area of the
greater U-shaped trough tends to curve lateroposteri-
orly around the anterior margin of the median ocellus
in E. liopeltis (as in fig. 139).
Like males, females of E. liopeltis and E. viperinus
can show striking similarities, but greater morpholog-
ical differences manifest themselves in structures of
the head and vulvar laminae. The trough anterior to
the median ocellus of E. /iopeltis is unlike the antero-
211
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
lateral pits of £. viperinus (fig. 137). The occiput of E.
viperinus (fig. 155) is much broader, and the anterior
margin of the occiput and curvilinear occipital crest
are roughly parallel. The median postoccipital swell-
ing of E. liopeltis is absent in E. viperinus. The vulvar
laminae of the two species are similar, but the Y-
shaped postlamellar ridge generally arises at or just
before the posterolateral margins of the lamellar
plates in £. /opeltis, the postlamellar ridge usually sur-
passes the vulvar lamina in £. viperinus. The median :
cleft between the vulvar lamellae is more obtuse in £.
viperinus (fig. 184) than in E. liopeltis (fig. 182).
Erpetogomphus liopeltis and E. viperinus are allopat-
ric and E. viperinus generally has more dense vena-
tion, despite wing lengths (£. /iopeltis hind wing 29-
31 mm [n = 13 males, 3 females], £. viperinus hind
wing 29-31 mm [n = 20 males, 20 females]). The
number of marginal cells behind fore wing paranal
cells in male £. liopeltis ranges from 0-2 (only one
wing has 3), compared with 1 (in 4 wings only) to 4
in È. viperinus.
Both sexes of E. liopeltisand E. elaps are easily diag-
nosed in the key by differences in body colouration,
shape, and position of the posterior hamular tooth
and shape of the cercus (males) and shape of the vul-
var laminae (females). The occiput of females of E.
elaps, E. elaphe, and E. liopeltis are similar in being re-
duced to a perpendicularly raised medial area (figs.
152-153) and a medial tumid area on the postocci-
put.
Remarks
Variation. — I found little variation among the 12
paratype males and 2 paratype females. The hind
margin of the occiput (crest) is medially concave in
the holotype (fig. 140), but ranges to almost linear
(fig. 141) in other specimens. A poorly defined nar-
row green stripe is present between the coalesced dark
antehumeral and humeral thoracic stripes in 7 males,
but this condition varies from a short, narrow mark to
one occupying the medial 0.50 of the lateral margin
of the mesepisternum.
Wing variation among paratypes: Males: anx: fore
wing 13-15/13-16, hind wing 9-12/10-12; pnx: fore
wing 8-10/8-10, hind wing 9-13/9-12; cs under pte-
rostigma: fore wing 5-7/4-6, hind wing 4-6/5-7; anal
triangular cells: 3-4/3-4. Hind wing 29-31 mm.
Abdomen 37-40 mm.
Females: Anx fore wing 16/14-15, hind wing
10/10; pnx fore wing 8-10/9-10, hind wing 11/9-11;
cs under pterostigma fore wing 6-7/6, hind wing 6-
717-8.
Other aspects of wing venation variability are char-
acterized under the diagnosis for E. hopeltisand E. vi-
perinus.
Some (n = 5) males have no marginal cells behind
212
fore wing paranal cells, while two others (fig. 227)
have one marginal cell in only one wing. All females
have at least two such cells, except for the left fore
wing of the Chipingue female. The fifth antenodal is
almost always thickened; but the sixth is thickened in
two males from La Estanzuela and in the right fore
wing of the Apatzingan female.
One male from La Estanzuela has aberrant hind
wing venation and was not included in the measured
samples. Most specimens, including the holotype,
had been preserved in acetone; and their resultant
preservation leads me to believe that the pale coloura-
tion is similar to the blue green of E. viperinus.
Biology. — Novelo (pers. comm.) misidentified this
species as E. elaps in Novelo and Pena (1991). They
collected E. hopeltis in Hidalgo state from 22 April to
25 July (Rio Zacuala, Pemuxtitla) and 27 July
(Calnali). They were taken as they perched on rocks
along narrow, shaded, shallow, rocky streams.
Erpetogomphus liopeltis was collected at Cerro El
Mirador with £. elaps. Data for the female collected
in Michoacan indicate that it was collected on ‘side of
stream’. Collection dates range from 22 April (Rio
Zacuala) to 17 August (Arroyo Dolores).
Distribution (fig. 201). — Erpetogomphus liopeltis is
only known from the states of Hidalgo, Michoacan,
Nuevo Leon, and San Luis Potosi in northeastern
Mexico.
Erpetogomphus bothrops sp. n.
(figs. 15-thx, 37-abd, 58-hamules, 80-penis, 102-
app, 118-cercus, 127, 128-ept, 136-base of
postfrons, 154-vertex, 183-vl, 201-distr, 228-wings)
Erpetogomphus viperinus. — Calvert 1899: 385 (described as
E. viperinus Selys); Calvert 1905: 163 (described as E. vi-
perinus Selys); Calvert 1909: 35 (misidentified as E. vi-
perinus, compared with E. schaust); Muttkowski 1910: 87
(in part, cat.); Williamson & Williamson 1930: 11 (in
part, refers to true E. viperinus but specimens in E. B.
Williamson Coll. misidentified); Calvert 1947: 608 (stat-
us of Tepic specimens); Paulson 1982: 256 (in part, as £.
viperinus from Mex., Guat.); Davies and Tobin 1985: 28
(in part, cat.); Tsuda, 1986: 87 (in part, cat.); Maes et al.
1988: 36 (as E. viperinus from Nicaragua); Tsuda 1991:
95 (in part, cat.).
Type material. — Holotype male: Mexico:
Veracruz: Rio Otapa, 8 km S of La Tinaja, elev. 90 m,
13 Aug. 1976 (R. W. Garrison). Allotype female:
same data, but 20 Aug. 1976. In UsNM.
Paratypes examined (614, 289). — Mexico:
Guerrero: Dos Arroyos, 1000 ft., Sept. 1888 (H. H.
Smith), 12 (BMNH); Jalisco: San Diego Rancho near
Cocula, 14 Nov. 1923 (J. H. Williamson), 16
(BMNH); Michoacan: El Sabino, 20 mi SSE of
Uruapan, 27 July, 1 Aug. 1936 (H. Devlin Thomas),
83, 32 (ummz, esca); Morelos: Puente de Ixtla, 3
July 1900 (C.C. Deam), 19 (ummz); Cerro del
Higuerón-Jojutla, July 1983 (C. Deloya), 1d
(UNAM); Nayarit. Acaponeta, 1-2 Nov. 1923 (J. H.
Williamson), 96, 12 (ummz); Tepic, 7 Nov. 1923
J. H. Williamson), 16 (ummz); Jumatän, 10 Sept.
1980 (G. Jiménez), 26 (UNAM); San Luis Potosi,
Cascadas Micos nr. aqueduct, 27 June 1990 (K. J.
Tennessen), 1d (KJT); El Salto, 400 m, 6 Sept. 1963
(T. W. Donnelly), 18, 19 (twp); Huichihuayan,
km 410, S of Valles, “Sam Brown’ Hacienda, 25 Sept.
1938 (L. J. Lipovsky), 16, 29 (ummz); Tamaulipas.
Rio Corona, 20 mi N of Ciudad Victoria, nr. Mex.
Hwy 101, 1000 ft., 26 July 1968 (R. W. Garrison),
Id, 22 (rwe); Veracruz Atoyac, 400 m,
(Schumann), 1 ® (BMNH); 16.5 mi S of Catemaco, by
Hwy 180, 25 June 1985, (I. S. Askevold), 24 (cc);
Cordoba 12-25 July 1964 (E. Fisher, D. Verity), 19
(LACM); Isla, 17 July 1969 (R. Wind), 14 (cc);
K375-390 Cordoba Rd., 11 Aug. 1961 (R. and K.
Dreisbach), 1d (usnm); 4.9 km N of Coscomatepec,
by Mex. Hwy 156, 11 Aug. 1976 (R. W. and J. A.
Garrison), 12 (Rwe); Rio Otapa, 8 km S of La
Tinaja, 90 m, 13, 20 Aug. 1976 (R. W. and J. A.
Garrison), 106, 22 (rwe); Rio Hondo, on road to
‘Colonia la Apachital’ (50°), Soteapan, 500 m, July-
Aug. 1990, (collector unknown), 6d, 4? (PM, RwG);
6 mi E of Hwy from Tierra Blanca to Ciudad
Aleman, 25-26 Aug. 1957 (G. H. Beatty, III), 1d
(rsca); Salto Eyipantla, 8 km S of San Andres Tuxtla
and Mex. Hwy 180, 15 Aug. 1976 (R. W. and J. A.
Garrison), 26 (RwG); Tierra Colorada, nr. Veracruz,
17 July 1932 (H. M. Smith), 19 (ummz); Oaxaca:
Candelaria Loxicha, 500 m, 7 Sept. 1973, (E. C.
Welling-M.), 1d (cc); 5 July 1974, (E. C. Welling-
M.), 1d (swp); Chiapas river 26.1 mi NE
Tapanatepec, 2100 ft., 2 Aug. 1965 (D. R. Paulson),
1d (pre); stream 15.4 mi NE Arriaga on Mex. 195,
2300 ft., 24 July 1965 (D. R. Paulson), 26 (pre); El
Aguacero, nr. Ocozocoautla, 26 Oct. 1986 (E.
Fisher), 16 (Rwa); GUATEMALA: Baja Vera Paz Dept.:
San Geronimo, 1879-80 (G. C. Champion), 1d
(BMNH); Chiquimula Dept: streams vic. Tierra
Colorado, 800 m, 20 July 1962, (T. W. Donnelly),
14,29 (twp); small stream vic. Veguitas, 600 m, 10
July 1962 (T. W. Donnelly), 16, 19 (rwp); Dept.
Zacapa: Gulan, 16-17 June 1909 (E. B. Williamson),
2d (ummz); La Union, 850 m, 31 Aug. 1972 (E. C.
Welling M.), 19 (ummz); EL SALVADOR:
Anuachopan Dept.: (all collected by V. Hellebuyck):
Bosque El Imposible, San Francisco Mendez, 16 Aug.
1987, 36 (va, RwG); May 1987, 1d, 19 (vm); 12
July 1987, 26 (vu); 25 July 1987, 16, 12 (vn); 16
Aug. 1987, 1d (vn); 3 Sept. 1987, 3d, 1 (1 pair in
copula) (vH); El Coyolar, El Imposible, 2 Aug. 1987,
26 (vx); Valle de la Puerta, Bosque El Imposible, 3
Sept. 1987, 3d (vx).
GARRISON: Revision of Erpetogomphus
Description
Holotype male. — Labium grey-white with wash of
brown around margin of median lobe; entire face in-
cluding vertex and occiput yellow green, a transverse
trough in front of median ocellus, lateral ends slight-
ly deeper than medial area; a small tubercle behind
each ocellus; pedicel, scape, flagellum brown; occiput
yellow green, mostly planar, slightly tumid medially,
crest green, slightly prominent, straight, slightly
notched medially and covered with long brown hairs;
postocciput green, transverse when viewed dorsopos-
teriorly; rear of head yellow brown, paler laterally; la-
teral margins of labrum and base of mandible ocher;
rear of head yellow brown.
Prothorax yellow green except for brown on anteri-
or margin of median lobe and posterior area of ante-
rior lobe. Synthorax (fig. 15) entirely yellow green ex-
cept for following brown areas: slight vague stripe
lateral to middorsal carina; antehumeral stripe; spot
on upper end of humeral suture connected anteriorly
to antehumeral, and gradually disappearing basally
toward mesinfraepisternum; line bordering antealar
crest; metasternum pale yellow green. Coxae, femora
pale yellow green, becoming dark brown dorsodistal-
ly; tibiae, tarsi and armature black.
Wings hyaline, anterior margin of costa yellow, re-
mainder of venation black, pterostigma brown.
Venational details. Fifth antenodal thickened in all
wings; number of marginal cells behind fore wing
paranal cells: 1/3; anx: fore wing 15/15, hind wing
11/11; pnx: fore wing 11/13, hind wing 13/13; cs
under pterostigma: fore wing 6/6; hind wing 7/6;
anal triangular cells: 4. Hind wing 27 mm.
Abdomen (fig. 37) with segment 1 yellow green
with a wash of brown dorsally; segment 2 yellow
green with diffuse brown spot posterodorsally to auri-
cle; segment 3 broadly light green dorsally, interrupt-
ed laterally by a longitudinal brown stripe flaring at
transverse carina, inferior margin of tergite ivory; seg-
ments 4-6 similar to segment 3, but lateral brown
stripes progressively more extensive and each meeting
its neighbor dorsally at posterior end of segment; seg-
ment 7 with anterior 0.66 pale green white except for
black transverse carina, posterior 0.30 of segment red
brown; segments 8-10 red brown, darker dorsally,
carinae and lateral foliations black; posterior medial
margin of 9 pointed. Abdomen 35 mm.
Cercus (fig. 102) pale ocher, gently decumbent at
posterior 0.30, gradually narrowing toward end, tip
with a shiny black tooth; epiproct brown, gently
curved as is characteristic for the genus, tip of epi-
proct when viewed posteriorly (fig. 128) bidentate.
Accessory genitalia. Anterior hamule dark brown,
branched at distal 0.30, superior branch larger than
inferior branch; posterior hamule (fig. 58) pale,
roughly triangular with an anterior shoulder, tip with
215
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
a black tooth twisted laterally from broad axis of ha-
mule; penis with cornuae pointed but each with me-
dian shoulder as shown in fig. 80; lateral lobe promi-
nent, knife- like, serrated posteriorly.
Allotype female. — Similar to male with following
differences: synthorax with definite dark triangular
middorsal stripe, its base not touching collar; broad,
diffuse brown dorsolateral stripe on abdominal seg-
ment 2, brown lateral stripes on segments 3-6 with
their anterior ends ending at basal 0.20 of each seg-
ment so that lateral white connects with dorsal yel-
low-white middorsal stripe; cercus pale.
Vulvar lamina (figs. 183) small, cleft about as large
as each plate; each plate connected basally, each with
a raised shelf occupying anterior 0.5; postlamellar
ridge with Y-shaped juncture posterior to hind mar-
gin of lamina; central stem with a well defined circu-
lar or oval depression on each side. Abdomen 32 mm.
Venational details. Fifth antenodal thickened in all
wings; no marginal cells behind fore wing paranal
cells; anx: fore wing 12/13, hind wing 9/9; pnx: fore
wing 8/9, hind wing 10/10; cs under pterostigma:
fore wing 4/5, hind wing 5/5. Hind wing 27 mm.
Diagnosis
Selys’ (1868) description of Erpetogomphus viperi-
nus was too brief and inexact for Calvert (1899) to de-
tect a specific difference between his specimens of E.
viperinus (= E. bothrops) and the true £. viperinus.
Examination of the lectotype male and female of £.
viperinus shows that the Erpetogomphus viperinus of
Calvert and all subsequent authors is referable to the
new species £. bothrops.
E. bothrops is most similar to E. liopeltis and is diag-
nosed under that species. Erpetogomphus bothrops also
resembles E. viperinus, but in life, the pale coloura-
tion of E. bothrops is yellow green (deep green in E.
viperinus). Erpetogomphus viperinus is generally larger
(hind wing male 29-31 mm) and darker. In males,
the cercus of £. bothrops is pale with a distinct black
tooth; while in E. viperinus this structure is dark red
brown with no apical tooth. In E. bothrops, the distal
0.5 of the cercus gradually narrows, so the distal 0.5
of the cercus in lateral view is not as thick or robust as
the base (fig. 102). In E. viperinus, the distal 0.5 of
the appendage is robust and is as thick as its base (fig.
103). In dorsoposterior view, the posterior 0.30 of
the cercus of E. bothrops gradually narrows (fig. 118),
and is not as laterally compressed as in E. viperinus
(fig. 119). The inferior distal margin of the cercus of
E. viperinus is strongly carinate, not so in E. bothrops.
The tip of each epiproct of E. bothrops in posterior
view is bidentate (fig. 128) or truncate (fig. 127), not
bluntly pointed as in E. viperinus (fig. 129). Other
characters are given in the key.
Females of these species are easily distinguished by
214
overall body colouration in life, and thoracic macula-
tion. Two structural characters separate E. bothrops
from E. viperinus. the anterior margin of the vertex of
E. viperinus contains two well defined pits, each dor-
solateral to the median ocellus. Each pit is the recep-
tacle for the bluntly pointed tip of the male epiproct
when in copulation. In E. bothrops, these pits are re-
placed by a continuous groove deepest medially,
which accommodates the explanate tips of the male
epiprocts. Secondly, the V-shaped notch of the vulvar
lamina in £. bothrops is more acute (fig. 183) than the
more widely divergent notch of E. viperinus (fig.
184). The shape of the vulvar lamina of E. bothrops
(fig. 183) distinguishes it from similarly marked fe-
males of E. sipedon (figs. 187, 188) and E. elaps (fig.
181).
Remarks
Variation. — Venational variation among the para-
type series of males (fig. 228): number of marginal
cells behind fore wing paranal cells: 0-3; anx: fore
wing 14-16/13-16, hind wing 9-12/10-12; pnx: fore
wing 9-11/8-13, hind wing 10-13/10-13; cs under
pterostigma: fore wing 5-7/5-7, hind wing 5-7/5-7;
number anal triangular cells: 3-4. Hind wing 21-28
mm.
Variation exists in the direction of the apical tooth
of the posterior hamule. It is present in all males and,
in most, is twisted so that its tip is directed laterally
(ge, distally) to the direction of the planar surface of
the hamule. In a few males, the tooth, though rotat-
ed, points in a more lateroanterior direction.
The tip of the epiproct, when viewed posteriorly, is
planar and bidentate (e.g., fig. 128); but in some, the
distal and mesal teeth are reduced. In rare cases, the
tip is almost straight (fig. 127); variations link these
extremes. I have found no correlation of the posterior
hamular condition with the epiproct condition; nor
do these conditions seem to vary according to altitude
or locality. For example, of 5 males from Nayarit,
Jamatan, only one has hamular teeth pointing pre-
dominantly cephalad. This same specimen has a bi-
dentate condition of tips of the epiprocts. Another
male has a bidentate condition, another has an inter-
mediate condition, and two others have truncate tips.
Of another series of 9 males (including holotype)
from the Veracruz, Rio Otapa, two have the hamular
tooth pointing predominantly cephalad. These two
specimens have bidentate epiprocts, although the
right epiproct of one male is intermediate. The re-
maining 7 males have bidentate epiprocts, but one
has an intermediate condition of its right epiproct.
The membranous hood of the penis is variable
among and within populations. Most specimens, in-
cluding the holotype, possess a short hood, exposing a
subcutaneous membrane (fig. 80); but a male from
San Luis Potosi, Cascadas Micos, has a long, acute,
non-overlapping hood. Another two males, one from
Guatemala (Dept. Chiquimula, small stream, vic.
Veguitas), and all specimens I examined from El
Salvador (Dept. Anuachopan, Bosque El Imposible,
San Francisco Mendez), have long, overlapping hood
membranes similar to that illustrated for the holotype
male of E. leptophis (fig. 76). With the exception of the
San Luis Potosi male, specimens with long, acuminate
hoods seem to typify most southerly specimens. I can
find no other characters differentiating these speci-
mens from others.
Venational variation among the paratype series of
females (n = 18): number of marginal cells behind fore
wing paranal cells: 1-3; anx: fore wing 12-16/12-16,
hind wing 9-11/8-11; pnx fore wing 8-12/9-12, hind
wing 9-13/9-13; cs under pterostigma: fore wing 4-
715-7, hind wing 5-8/5-7. Hind wing 27-31 mm.
Biology. — I have often seen this species along mar-
gins of cut agricultural fields bordering streams and
canals. Its apple green colouration renders it difficult
to detect among vegetation, but when disturbed, it
does not fly far. The type locality is a wide, shallow
stream bordered by trees in extensively cultivated
farmland. I collected E. bothrops next to cut fields
along with six other gomphids: Erpetogomphus eutai-
nia, E. ophibolus, Phyllocycla breviphylla Belle, P. vol-
sella (Calvert), Phyllogomphoides — duodentatus
Donnelly, and Progomphus clendoni Calvert. Twenty-
five other species of Odonata were collected there.
R. Novelo (pers. comm.) observed a pair in copula
in Morelos state (Cerro del Higuerón) at 1250 m at
18.20 hr on 14 Sept. 1988.
Williamson records the following on two
Acaponeta males: ‘Easlilly caught while sitting in
brushy weed patch on bank of river, sun behind
cloud,’ and ‘Quite common flying over water just
above the real rapids and lighting on willow-like stems
and twigs along the bank. Saw no ® gomphine today.’
López (in litt.) writes that they perch along sides of
streams or on exposed rocks in the middle of streams.
He states that they do not remain long at a spot, mov-
ing constantly along the length of the stream.
Elevation gradients range from near sea level (50
m, Veracruz, Agua Caliente) to 1250 m (Morelos,
Cerro del Higuerén). Collection dates range from
May (El Salvador) to November (Mexico, Jalisco).
Distribution (fig. 201). — This species and E. elaps
are the most widely distributed species in Mexico and
Guatemala. Ærpetogomphus bothrops ranges from
Tamaulipas in eastern Mexico (24°N) south to El
Salvador (14°N). It also occurs in mesic areas in west-
ern Mexico from Nayarit (22°N) south. Its distribu-
tion indicates an avoidance of the xeric areas in north-
western Mexico and the high mountain plateau in
northern Mexico.
GARRISON: Revision of Erpetogomphus
Erpetogomphus viperinus Selys
(figs. 16-thx, 38-abd, 59-hamules, 81-penis, 103-
app, 119-cercus, 129-pet, 137-base of postfrons,
155-vertex, 166-postocciput, 184-vl, 201-distr, 229-
wings)
Erpetogomphus viperinus Selys, 1868: 68 (3 sep.) (descr. of
3, 2, ‘D’Orizaba’). — Selys 1869: 176 (13 sep.) (rede-
scription of d and 9 from Orizaba); Selys 1873b: 519
(75 sep.) (list); Muttkowski 1910: 87 (in part, cat.);
Williamson & Williamson 1930: 14 (summary of status);
Paulson 1982: 256 (Mex.); Davies and Tobin 1985: 28
(cat.); Tsuda 1986: 87 (cat.); Bridges 1991: VII.220
(cat.); Tsuda 1991: 95 (cat.).
Herpetogomphus viperinus. — Hagen 1875a: 42 (cat.); Selys
1879: 64 (2 sep.) (status of classification); Kirby 1890: 60
(cat.).
Description
Male. — Entire head pale green (blue green in life),
slightly darker on anterior of ante- and postclypeus
and labrum, lateral margins of labrum ocher, base of
mandible and labium dull grey green; basal 0.25 of
postfrons and all of vertex brown, a pair of pits at an-
terior margin of vertex at juncture of base of antefrons
(fig. 137), one each anterolateral to median ocellus;
antennae dark brown; occiput with well developed
medial tumid area, green, occipital crest slightly
emarginate medially, dark brown to black, rimmed
with long dark brown hairs; postocciput brown, gr-
een medially, transverse, with a slight vertical depres-
sion medially; rear of head yellow brown.
Prothorax brown, anterior and posterior lobes
green. Synthorax (fig. 16) entirely green (blue green
in life) with following dark brown: obscure middorsal
stripe ending before collar, becoming darkest at base
of antealar crest and extending as narrow wash of
brown below rim of antealar crest and joining wide
stripe along lateral 0.30 of mesepisternum; this stripe
coalescing with normal humeral stripe; often with a
narrow isolated stripe of green separating these two
stripes; combination antehumeral and humeral stripe
reaching collar; all of mesinfraepisternum and ventral
part of thorax; rim below subalar carina, often with
small extension on obsolete second lateral suture,
which in some specimens may form an indistinct sec-
ond lateral stripe; narrow but well defined third later-
al stripe. Metasternum pale grey green. Coxae, tro-
chanters grey, becoming brown exteriorly; base of
femora brown, becoming black distally; tibiae and ar-
mature black.
Wings (fig. 229) hyaline, anterior margin of costa
dark brown at base, becoming yellow distally to prox-
imal level of pterostigma, remainder of venation and
pterostigma black.
Venational statistics. Fifth (occasionally sixth) an-
tenodal thickened in all wings; number of marginal
cells behind fore wing paranal cells: 1-4/1-4; anx: fore
AMS)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
wing 13-18/13-18, hind wing 9-12/10-13; pnx: fore
wing 8-13/10-12, hind wing 10-14/10-13; cs under
pterostigma: fore wing 5-7/4-7, hind wing 5-7/5-7;
anal triangular cells: 4 (rarely 5). Hind wing 29-31
mm.
Abdomen (fig. 38) with segment 1 green with
brown dorsolaterally except for green on posterior
dorsal 0.50 of segment; segment 2 with middorsal
green stripe; brown dorsolaterally surrounding green
auricle; ventrolateral area pale green, annulus black;
segments 3-6 primarily black with following pale are-
as: basal 0.10 of each segment, dorsally forming an in-
complete blue green middorsal stripe extending to
posterior 0.20 of each segment; ventrolateral wedge
shaped white spot at middle of each segment, con-
necting in some specimens to basal pale ring; segment
7 with dorsal 0.50 light green, posterior 0.50 red
brown, black along transverse carina; segments 8-10
predominantly red brown, darker dorsally, denticu-
lated posterior margin of these segments black; poste-
rior margin of segment 9 often forming a posteriorly
directed point. Abdomen 36-40 mm.
Cercus (fig. 103) pale ocher, slightly darker ven-
trally, robust and strongly curved at posterior 0.30,
inner margin of posterior 0.30 of cercus planar (fig.
119); same structure in posterodorsal view narrow,
inferior carina along posterior 0.30 of cercus; epiproct
brown, becoming black posteriorly, gently curved as
is characteristic for the genus; tip of epiproct, when
viewed posteriorly (fig. 129) forming a narrow, blunt
point.
Accessory genitalia. Anterior hamule (fig. 59)
black, divided at distal 0.30; smaller posterior branch
almost as large as anterior branch, its pointed tip al-
most meeting tip of anterior (larger) branch; posteri-
or hamule (fig. 59) pale, roughly triangular, swollen
at distal 0.25 to 0.50 of appendage, tip with a black
tooth twisted laterad from broad axis of hamule; penis
with cornuae pointed but each with well developed
median shoulder (or median lobe in some specimens),
as shown in fig. 59; lateral lobe prominent, knife-like,
arcuate laterally, its outer margin strongly serrated.
Female. — Similar to male, with following differ-
ences: Vertex (fig. 155) without postocellar ridge, a
well defined pit anterolaterad to median ocellus as in
male; occiput green, narrow, mostly planar, or with
only a slight tumid area medially, hind margin slight-
ly sinuate, barely emarginate at middle; postocciput
green, transverse; synthorax with no vestige of second
lateral stripe; abdomen with dorsolateral dark stripe
on segment 1 reduced or vestigial, middorsal dark
stripe on segments 2-6 narrower, anteriorly touching
black annulus, widened at transverse carina, con-
stricted near center and widening at distal 0.25 of
each segment, ventral white narrow but expanded
near center of segment, sometimes a longitudinal
216
wedge-shaped spot isolated by narrow posterior and
anterior lateral offshoots of black dorsolateral stripe,
white lateral wedge- shaped spots largest on segments
2 and 3, dividing or partially dividing dorsolateral
brown stripe; segment 7 similar to segment 6, but
dorsal pale orange green connecting anteriorly with
lateral white at basal 0.10 of segment; segments 8-10
dark brown, becoming black dorsally and posterolat-
. erally to all black with ill-defined white lateral spot on
each segment; cercus, paraproct brown. Abdomen
37-41 mm.
Vulvar lamina (fig. 184) small, cleft as large as each
plate; each plate broadly connected basally, each with
a well developed raised shelf occupying anterior 0.50;
medial margin of each plate slightly concave, its tip
falcate; postlamellar ridge with Y-suture posterior to
hind margin of lamina; central stem short with a well-
defined circular or oval depression on each side.
Venational statistics. Anx: fore wing 15-20/15-21,
hind wing 10-14/11-14; pnx: fore wing 10-13/10-15,
hind wing 10-15/9-14; cs under pterostigma: fore
wing 5-8/5-8, hind wing 5-8/6-7. Hind wing 31-34
mm.
Diagnosis
Erpetogomphus viperinus is most closely related to
E. bothrops, and is diagnosed under that species. It is
also superficially similar to £. liopeltis and is diagno-
sed under that species. The thoracic pattern of £. vi-
perinus is superficially similar to that of £. sipedon, but
is easily separated by characters listed under that spe-
cies and in the key.
Remarks
Variation. — The description is based on 49 males and
38 females from Veracruz State, Mexico. The lectotype
and paralectotype are in reasonably good condition, but
post mortem preservation is poor and would yield an in-
adequate description. Venation characters for this pair
are as follows:
Lectotype male: number of marginal cells behind fore
wing paranal cells: 3/1; anx: fore wing 16/17, hind wing
12/12; pnx: fore wing 13/12, hind wing 11/13; cs under
pterostigma: fore wing 6/6, hind wing 6/5; anal triangu-
lar cells: 4/4.
Paralectotype female: number of marginal cells be-
hind fore wing paranal cells: 4/3; anx: fore wing 15/18,
hind wing 12/11; pnx: fore wing 10/12, hind wing
12/13; cs under pterostigma: fore wing 7/8; hind wing
716.
Examination of 20 males shows variability is some ve-
national characters. All specimens have marginal cells
behind the fore wing paranal cells, ranging from 1-4
cells. The fifth antenodal is most often thickened, but
rarely the fourth (one wing) or sixth (7 wings). The sixth
antenodal is thickened on both wings of the lectotype.
Little variation exists in body colouration. The sec-
ond lateral thoracic stripe is incomplete or vestigial in
most males. It is complete on only one male from Rio
Metlac. Females show a greater tendency toward a
dense venation than do males. Marginal cells behind
the fore wing paranal cells were present in all 20 fe-
males I examined, ranging from 1 (one wing) to 8
(two wings). The fifth antenodal is generally thick-
ened, but it is the fourth on the left wing of the para-
lectotype, sixth in 17 wings (including right wing of
paralectotype) and seventh in four wings.
Biology. — Although of restricted distribution, £.
viperinus can be common during certain years. I col-
lected a few specimens of both sexes in the mostly
shaded understory at the Rio Metlac. The locality
consists of a fast running river in a steep canyon. The
narrow valley contained many coffee plants. I flushed
various specimens, and they alit on low vegetation,
where their deep blue green colour made them diffi-
cult to detect unless the eye followed closely where
each specimen landed. Once approached they were
easy to take with a net. Other Odonata taken at the
same site were Hetaerina cruentata Rambur, Argia sp.
nr. fissa Selys, Argia extranea Hagen in Selys, Aeshna
psilus Calvert, Brechmorhoga pertinax (Hagen), B. vi-
vax Calvert, Cannaphila vibex Hagen, Erythrodiplax
fusca (Rambur), and £. umbrata (Linnaeus).
The species has been collected more frequently far-
ther north at Teocelo under similar circumstances.
The Odonata assemblage there is more varied and
consists of Palaemnema sp. n., Paraphlebia zoe Selys,
Argia extranea, Argia sp. n., Brechmorhoga pertinax, B.
rapax Calvert, B. tepeaca Calvert, Cannaphila vibex,
Libellula herculea Karsch.
This species has been taken at elevations of 800-
1300 m (Teocelo and vicinity). Collection dates
range from 6 June through 18 September (Teocelo
and vicinity).
I have abstracted the following biological notes
from López (in litt): Tenerals and adults travel far
from the stream, hiding in vegetation in open areas,
where they feed by making short sallies from perches.
They capture primarily small flies and butterflies
(Lycaenidae) passing by. Copulation takes place in
open fields away from streams, as males grab females
which fly by. The copulating pair then retires to the
dense forest. Females oviposit while flying above wa-
ter, dropping their eggs from about 30 cm. At dusk,
males return to feeding sites, often in considerable
numbers, where threat display patterns among males
and females have been observed when two or more
individuals try to occupy the same perch. Specimens
are often preyed upon by spiders (Araneidae).
Distribution (fig. 201). — Erpetogomphus viperinus
is apparently restricted to central Veracruz. As stated
under the species account for E. bothrops, all pub-
GARRISON: Revision of Erpetogomphus
lished records of this species since Calvert’s redescrip-
tion of E. viperinus refer to the widespread £. both-
rops. I examined one male from the Paris Museum
with a handwritten label, ‘N. Carolina’, but I have
seen no specimens of E. viperinus from the United
States. I believe that locality is in error.
Material
Type data. — Lectotype male by present designa-
tion with following data: small green handwritten la-
bel ‘Mex/B[ouchard]’; white handwritten label “Herp.
/ viperinus /S.{elys]/ 3°; two manila coloured labels
each with ‘26’ handwritten in pencil in an unknown
hand; rectangular red label with printed ‘LECTOTY-
PE /and handwritten: “Erpetogomphusl viperinus Selys
1869/ G /des. [printed] R. W. Garrison 1984. The
caudal appendages are shown in fig. 103.
Paralectotype female: small green handwritten label
‘Mex./Blouchard]’; white handwritten label “Herp./
viperinus! S[elys]! 2’, two manila coloured labels,
each with ‘28’ handwritten in pencil in an unknown
hand; rectangular white label with printed ‘LECTOTY-
PE / and handwritten ‘Erpetogomphusl viperinus Selys
1869/ 9 / des. [printed] R. W. Garrison 1984. Both
specimens in IRSN.
Other material (485, 329, including lectotype d
and paralectotype 2). — Mexico: Veracruz. nr.
Municipio Teocelo at Puente Teocelo, ca. 1150 m,
(UNAM, RWG); Barranca de Cayoapa, Teocelo, 750-
890 m, (UNAM, JB, CE, PSM, DALD); Barranca de
Cayoapa, Tejeria-Teocelo, 800 m, (UNAM); road to
Monte Blanco, Teocelo, (UNAM); El Trapiche,
Teocelo, 1100 m (UNAM); road to Santa Rosa,
Teocelo, (UNAM); Santa Rosa, Teocelo, (UNAM);
Cascada de Xico, Teocelo, 1100 m, (UNAM); Cascada
Texolo, (UNAM); Parque Javier Clavijero, Jalapa,
1300 m (UNAM); Fortin de las Flores, 1010 m (RwG);
Rio Metlac, ca. 3.5 km WNW of Fortín de las Flores,
900 m (RwG); km 327 W of Cordoba, (rsca);
Cordoba, (USNM).
Erpetogomphus designatus Hagen in Selys
(figs. 17, 18-thx, 60-hamules, 82-penis, 104-app,
130, 131-ept, 156, 157-vertex, 185, 186-vl, 202-
distr)
Erpetogomphus designatus Hagen in Selys, 1858: 661 (401
sep.) (descr. d, 2, “Pecos River, Texas’). — Hagen in Selys
1859: 536 (10 sep.) (descr. d, 2); Selys 1873b: 519 (75
sep.) (list); Calvert 1899: 386 (mentioned); Calvert 1905:
166 (Ohio, Ind., Mexico); Muttkowski 1910: 86 (cat.);
Calvert 1912a: 289 (Mex., distr.); Hine 1913: 96 (Ohio);
Williamson 1914b: 447 (Tex.); Kennedy 1917a: 544
(larva, notes); Williamson 1917: 8 (Ind.): Kennedy
1917b: 137 (Kans.); Kennedy 1918: 298 (notes);
Williamson 1923: 8 (Kentucky); Montgomery 1925:
386 (Ind., habits); Montgomery 1927: 289 (Ind.);
DAY
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Kennedy 1928: 373 (seasonal distribution); Montgomery
1929: 340 (Ind.); Needham & Heywood 1929: 80 (key,
descr.); Byers 1930: 53 (Fla. Key, descr., habits);
Williamson & Williamson 1930: 12 (summary of status);
Byers 1931: 51 (Tenn.); Bird, 1932: 51 (Okla.);
Williamson 1932: 23 (Mo., habits); Tinkham 1934: 216
(Tex.); Montgomery 1935: 234 (Ind.); Borror 1935: 453
(Ohio); Borror 1937: 186 (Ohio); Montgomery 1937:
207 (Ind., habits); Ahrens 1938: 11 (Mo., predation);
Wright 1938: 27 (Tenn.); Ferguson 1940: 5 (Tex.);
Montgomery 1940: 289 (S. C.); La Rivers 1940b: 63
(Nev.); Montgomery 1941: 230, 238 (Ind.); Ferguson
1942: 146 (Tex.); Montgomery 1947: 165 (distr.); Bick
1951: 179 (Okla.); Montgomery 1951: 207 (Ind.);
Needham and Westfall 1955: 146 (key, descr.); Cross
1955: 11 (S. C.); Montgomery 1955: 133 (Ind.); Cross
1956: 4 (Fla.); Bick and Bick 1957: 2 (Okla.); Kormondy
1957: 108 (Tenn.); Bick 1957: 80 (La.); Gloyd 1958: 8
(Tex.); Bick and Bick 1958: 240 (Okla.); Bick 1959: 131
(Ark.); Donnelly 1961: 7 (Md. Va., D. C.); Alrutz 1961:
23 (Ohio); Borror 1963: 104 (common name); Macklin
and Cook 1967: 120 (Ky.); Montgomery 1967: 127
(distr.); Roback & Westfall 1967: 114 (water quality
data); Montgomery 1968: 134 (distr.); Pinhey 1969: 189
(tandem linkage); Resener 1970: 37 (Ky.); Huggins et al.
1976: 16 (Kans.); Harp & Rickett 1977: 50 (Ark.);
Huggins 1978: 2 (Kans.); Young & Bayer 1979: 90 (lar-
va, key, Tex.); Carle 1979: 322 (status in Virginia);
White et al. 1980: 26 (S. C.); Carle 1982: 339 (W.Va.);
Dunkle & Westfall 1982: 32 (status in Fla.); Paulson
1982: 255 (U.S., Mex.); Huggins & Brigham 1982: 4: 39
(N. C., S. C.); Harp & Rickett 1985: 132 (Ark.); Davies
& Tobin 1985: 27 (cat.); Tsuda 1986: 87 (cat., attributes
authorship to Selys); Bridges 1991: VII.60 (cat.); Tsuda
1991: 95 (cat.); Dunkle 1992: 39 (Fla).
Gomphus designatus. — Hagen 1861: 99 (descr. 5, 9).
Gomphus sp.. — Cabot,1872: 4 (descr. larva as Gomphus sp.
No. 6, Poles Creek, Tex.).
Herpetogomphus designatus. — Walsh 1862: 389 (compared
with Ophiogomphus rupinsulensis); Hagen 1875a: 42
(cat.); Selys 1879: 64 (2 sep.) (characters of genus);
Hagen 1885: 255 (larva of Cabot identified); Kirby 1890:
60 (cat., attributes authorship to Selys); Banks 1892: 179
(Kans.); Calvert 1899: 386 (characters of genus); Adams
1900: 622 (Ark.); Williamson 1903: 226 (Tenn.); Van
der Weele 1906: 177 (mentioned); Tucker 1907: 79
(Kans.); Tucker 1908: 99 (Tex.); La Rivers 1938: 76
(Nev., habits); La Rivers 1940a: 112 (Nev.).
Description
Male. — Entire face pale green, more vivid on post-
clypeus and frons, vertex with wash of brown around
ocelli or entirely dark brown, scape dark brown, pedi-
cel light brown, flagellum dark brown; anterior mar-
gin of vertex forming a trough at junction of ante-
frons, this trough with a pair of deeper pits
anterolateral to median ocellus; occiput green, wide,
tumid medially, crest slightly sinuate, barely emargi-
nate medially or straight, its hind margin covered
with long brown hairs; postocciput not visible dorsal-
ly, green; rear of head brown, darker toward occipital
foramen.
Prothorax predominantly brown, becoming green
218
dorsally; anterior and posterior lobes green; synthorax
(fig. 17) green with following dark brown: well de-
fined middorsal stripe widening to collar, its upper
end extending along antealar sinus connecting with
narrow humeral stripe; an isolated antehumeral stripe
not touching dorsal or ventral margin (except in spec-
imens from Durango and Chihuahua — see remarks);
lower part of humeral stripe extending posteriorly to
. form a fragmented second lateral stripe, though in
some specimens a vestige of a complete stripe exists;
narrow third lateral stripe. Coxae, trochanters pale
green, femora green becoming brown posterolaterally
at distal 0.20; distal 0.05 of metafemora green with
narrow brown line extending distally from subapical
brown area; tibiae dark brown with yellow along lat-
eral carinae of basal 0.5 of mesotibiae, a vestige of yel-
low or none on metatibiae; tarsi dark brown, arma-
ture black.
Wings hyaline with wash of yellow basally, vena-
tion dark brown, basal wing venation light brown, es-
pecially anteriorly, anterior margin of costa yellow,
except for anterior margin of pterostigma; pterostig-
ma dark brown, veins bordering it black.
Venational statistics. Fifth (rarely fourth) anteno-
dal thickened in all wings; number of marginal cells
behind fore wing paranal cells: 2-5/1-4; anx: fore
wing 11-14/11-14, hind wing 8-11/8-11; pnx: fore
wing 7-10/8-11, hind wing 7-11/8-12; cs under pte-
rostigma: fore wing 5-7/5-7, hind wing 4-7/4-7; anal
triangular cells: 3-5/3-4. Hind wing 28-31 mm.
Abdomen. Segment 1 green with dorsolateral
brown stripe occupying basal 0.75 of segment; seg-
ment 2 similar to segment 1, but dark brown expand-
ing ventrolaterally behind auricle, contracting again
and connecting with dark brown annulus; segment 3
similar to segment 2 but dark brown dorsolateral
stripe interrupted at anterior 0.25 and again at distal
0.50 to 0.75 of segment, dark brown prominent at
transverse carina, posterior 0.25 to 0.30 of segment
with dorsolateral brown expanding laterally to encir-
cle segment at posterior 0.10 of segment; segments 4-
6 similar to segment 3 but dorsolateral brown inter-
rupted only at anterior 0.25 to 0.30 of segment and
expanding abruptly at transverse carina; segment 7
with narrow dark brown on transverse carina and
poorly defined brown on lateral 0.50 of segment; seg-
ments 8-10 yellow brown with dorsolateral red
brown occupying basal 0.40 to 0.75 of segments; pale
areas of segments 2-6 orange green dorsally, becom-
ing white below. Abdomen 35-37 mm.
Cercus (fig. 104) yellow brown, strongly angulate
near middle; tip of cercus beyond angulation narrow,
drawn out into a fine point; ventral margin with infe-
rior carina at basal 0.45. Epiprocts yellow brown,
gently curved dorsally at 90°, tips obliquely truncate
(fig. 130) or spatulate (fig. 131).
Accessory genitalia (fig. 60). Anterior hamule
black, divided at upper 0.50; lower branch forming a
small, posteriorly directed truncate appendage; poste-
rior hamule pale green, triangular, with anterior basal
shoulder, tip evenly round, black; penis with long,
decumbent lanceolate lateral lobe, its posterior end
serrated; cornuae separated, tips narrowly rounded,
with no medial lobe; membranous hood moderately
long but usually with ends not overlapping.
Female. — Head as in male, but median swelling of
occiput more prominent; prothorax and synthorax as
in male; wings with yellow at base more extensive, of-
ten with markings at base of arculus, to cubitoanal
crossvein; abdomen with definite pale orange dorsal-
ly, white to orange white laterally, these areas separat-
ed by brown dorsolateral stripes as in male; these
stripes on each segment may be narrower, especially
so between transverse carina and posterior 0.20 to
0.30 of segment; segments 8-9 mostly dark brown
dorsally, sides tawny, annulus pale; segment 10 yel-
low brown, cercus ocher. Abdomen 33-36 mm.
Vulvar lamina (figs. 185-186) with prominent ba-
sal plates; quadrangular laminar plates acuminate
posteriorly, meeting at medioanterior margin; cleft
between plates broadly U-shaped; juncture of Y-
shaped postlamellar ridge at or before level of posteri-
or margin of lamina. Abdomen 33-36 mm.
Venational statistics: number of marginal cells be-
hind fore wing paranal cells: 2- 5/1-5; anx: fore wing
12-15/11-15, hind wing 8-12/8-11; pnx: fore wing 7-
12/7-12, hind wing 8-12/7-12; cs under pterostigma:
fore wing 5-7/5-7, hind wing 4-8/5-7. Hind wing 28-
32 mm.
Diagnosis
Little difficulty should be encountered in identi-
fying either sex of this species. The strongly angulate
cercus superficially allies it with £. sipedon, E. hetero-
don, and E. lampropeltis, but the structure of the occi-
put and the penile characters easily separate males of
these species. The broadly defined occiput, with the
large medial tumid area characteristic of E. designatus,
is not present in the other three species. Male E. lam-
propeltis have a broad occiput, as do male E. designa-
tus, but the medial area of the occiput is only slightly
tumid.
The long, recumbent lateral lobe of the penis is
unique to this species (fig. 82). In £. lampropeltis and
E. sipedon, this structure forms a spinulose semicircle
(figs. 83-85).
As in males, females of E. designatus are distin-
guished from all other similar species by the tumid
median area of the occiput (fig. 156). The vulvar lam-
ina in E. designatus (figs. 185- 186) more closely ap-
proaches that of £. bothrops (fig. 183) than of E. sipe-
don (figs. 187-188), E. heterodon (fig. 193), or E.
GARRISON: Revision of Erpetogomphus
lampropeltis (figs. 189-190). Each lamellar plate of £.
designatus is smaller and the notch in the medial mar-
gin is more concave than in the other species.
Remarks
Variation. — Calvert (1907) described a specimen
from San Pedro, Coahuila, Mexico, which differed fr-
om typical £. designatus in the following characters
(typical condition in parentheses): 1) dark antehu-
meral stripe reaching down to mesinfraepisternum
(lower part of stripe isolated), and 2) darker, more
pronounced markings on abdominal segments as fol-
lows: segment 7 with a black spot on each side of pos-
terior 0.50 of segment (like area red brown, blending
anteriorly with ocher pale area); segment 8 with a
black stripe on each side for almost its entire length
(this area red brown); segment 9-10 with black at ba-
sal dorsolateral 0.50 to 0.75 (this area red brown with
some black on dorsal 0.50 of segment). Calvert
(1907) considered the San Pedro male to represent £.
designatus. This specimen is indicated as in the ANSP,
but it could not be found (Azuma in litt, 1984).
However, I have seen three males (Durango and
Chihuahua) and a female (Durango) whose markings
agree with Calvert’s San Pedro male. I also find the
following other differences: 1) no wash of yellow pre-
sent on wing bases, 2) dark thoracic stripes thicker
and more pronounced (fig. 18), 3) a black spot anter-
oventrally to auricle of abdominal segment 2, and 4)
tip of epiproct, viewed posteriorly, is spatulate (fig.
131), not obliquely truncate (fig. 130) as is typical of
other specimens. The single female has a more arcu-
ate occiput (fig. 157) than typical E. designatus (fig.
156), but the dark anteroventral spot on segment 2 is
small and inconspicuous.
Specimens from Durango, Chihuahua, and
Coahuila could be a well-defined subspecies charac-
terized by overall darker colouration and slight mor-
phological modification of the female occiput (if typ-
ical for this form) and male epiproct. I detect no
morphological differences in the penes, hamules, or
cerci and, like Calvert, I am inclined to treat these
specimens as E. designatus. Further evidence to sup-
port their conspecificity comes from a male from
Nuevo Leon (in foothills of the northernmost range
of mountains) in which the antehumeral stripe ex-
tends to the mesinfraepisternum but is in all other
characters typical of E. designatus.
One male from McLennan Co., Texas, has the tips
of the posterior hamules with a small, anteriorly re-
curved hook. This male and another from the same
locality have vestiges of an anteroventral dark spot on
abdominal segment 2.
I have not seen sufficient material of this species
throughout eastern parts of its range to render any
conclusions about geographic variability. Numerous
219
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
specimens, primarily from Gonzales and Williamson
counties, Texas, show little variability.
Although the type locality for £. designatus and
many other species collected by Capt. John Pope of
the Smithsonian Institution and described by Hagen,
is said to be the Pecos River in western Texas (32°N,
104°W), Needham and Cockerell (1903), citing
Scudder and Cockerell (1902), argue that the type lo-
cality was probably at or near Roswell, Chaves Co.,
New Mexico, as follows (Needham and Cockerell
1903: 138-139): ‘In Proc. Davenport Acad. Sci. vol.
9 (1902), p. 51, it is inferred that Pope collected his
material at very different localities above the river,
since he obtained both Melanoplus bivittatus and M.
differentialis, which inhabit different life-zones.
However, in 1902 it was observed that these two
grasshoppers do actually occur together at Roswell. As
it is not very likely that their ranges overlap very
much, it thus seems likely that Pope’s collections, if
all from one place, were from the vicinity of the
present town of Roswell, rather than from Lat. 32°
(the present boundary between New Mexico and
Texas), as Hagen indicates. In all probability, howev-
er, the collections were made in several places.’
No year is indicated on the data of any of the type
material, nor have I found any reference to the year of
Capt. Pope’s expedition. However, a public display
on the recent human history of the caverns at the vis-
itor center at Carlsbad Caverns National Park, Eddy
Co., New Mexico, states that Pope’s expedition oc-
curred in 1854-1855.
Venational details of lectotype male: number of
marginal cells behind fore wing paranal cells: 1/2; ;
anx: fore wing 12/13, hind wing 10/10; pnx: fore
wing 8/9, hind wing 9/9; cs under pterostigma: fore
wing 6/6, hind wing 7/6; anal triangular cells: 3/4.
Biology. — The synonymy indicates that more is
known about this species than any other congener.
Erpetogomphus designatus seems to be a species of mes-
ic deciduous forests where there are clear streams and
rivers. Montgomery (1925) writes of its habits in
Indiana: “This species was common in a well-cultivat-
ed orchard near Vincennes on August 7, 1924, rest-
ing on the ground with wings spread out against the
soil but taking flight so readily that only three speci-
mens were taken in two hours although I saw 20 or
more. In Hamilton Co., Indiana, Montgomery
(1937) records it as … rather numerous at ripples in
the river, flying just above the surface of the water,
and alighting on rocks. However, it was exceedingly
difficult to capture...’. Williamson (1932) observed
this species flying back and forth near the surface over
ripples in Missouri. Others alighted on vegetation or
on tips of mullein heads. La Rivers (1938) described
the flight of E. designatus at Hiko Springs in the
Pahranagat Valley, Nevada, as ‘low, swift, and change
220
of position rapid...’ Ahrens (1938) describes the cap-
ture of a male along with a male Hagenius brevistylus
Selys near Lebanon, Missouri: “The H. brevistylus had
struck and seized the designatus with such force that
both were carried into the water. Both were dipped
from the river an instant after they struck the surface.’
Dunkle and Westfall (1982) assign E. designatus as
‘threatened’ in Florida, because it is thus far known
. only from Liberty and Gadsden counties in the pan-
handle. In 1975 and 1977, I found this species abun-
dant along cultivated stubble bordering trees next to
the Guadalupe River in Gonzales County, Texas. It
was more abundant than £. eutainia, and both species
were easily flushed from low or cut vegetation.
Members of both sexes did not fly far, and I could
easily photograph and capture specimens.
Collection dates range from 6 May in Texas
(Williamson, 1914b) to 5 October in Tennessee
(Wright, 1938).
The larva has been described by Cabot (1872),
Needham and Heywood (1929) and Needham and
Westfall (1955).
Distribution (fig. 202). — Erpetogomphus designatus
and £. compositus are the most boreal of the genus,
with £. designatus generally replacing the western £.
compositus in the eastern United States.
Erpetogomphus designatus occurs as far north as
Maryland and Washington, D. C., west through
West Virginia, Ohio, Indiana, Missouri, Kansas,
southeastern Colorado, eastern New Mexico, into
western Texas. Sidney Dunkle (in /itt., 1991) men-
tions one female from South Dakota (Fall River Co.,
9.7 mi SW Hot Springs, 29 July 1976, G. and ].
Bick, L. Hornuff), erroneously published as Stylurus
intricatus (Selys) (Bick et al., 1977: 151), which is the
northernmost record for this species. It occurs gener-
ally throughout the southeastern United States, but
does not penetrate peninsular Florida (Dunkle and
Westfall, 1982). Farthest western records include
Hiko Springs, Lincoln Co., Nevada (La Rivers, 1938)
and Arizona (Maricopa Co., Salt River at Corn Bluff
Campground, 15 mi NE of Mesa). Erpetogomphus de-
signatus is found as far south as Nuevo Leon (24-
26°N, 100°W) and west to Coahuila (Calvert, 1907),
Durango, and Chihuahua, Mexico.
It is partially sympatric with £. compositus in
Arizona, Nevada, New Mexico, and western Texas. I
collected it with E. eutainia in Gonzales Co., Texas.
Material
Type data. — Lectotype male by present designa-
tion with following data: ‘July/ 16’ [handwritten],
‘designatus [handwritten], ‘Hagen’ [printed], small
white label with printed ‘Type’ with red label glued to
lower edge with handwritten ‘1864’; red rectangular
label with printed ‘LECTOTYPE/ and written
‘Erpetogomphus designatusl Hagen d/ des. [printed]
R. W. Garrison 1984. Another d , a paralectotype,
has the handwritten labels, ‘Erp. designatus! 3/ Pecos
River’ [in Hagen’s hand] and ‘Pecos River/ Uhler
1860.’ in an unknown hand. A female paralectotype,
collected ‘July/ 15° has the additional label: [all prin-
ted unless otherwise stated] ‘ERPETOGOMPHUS/ DE-
SIGNATUS HAGEN [stamped] / P. P. Calvert, det [writ-
ten] 1905/ B. C. A. Neur., p. [written] 166.’ Type
locality herein restricted to vicinity of Roswell,
Chaves Co., New Mexico (see remarks).
Other material (844, 532, including lectotype d
and d ® paralectotypes). — U. S. A: FLoripa:
Gadsden Co., Apalachicola River, Aspalaga Landing,
Hwy 1-10 (jp, Rwe); GEORGIA: Houston Co.,
Ocmulgee River at 96, (RWG); VIRGINIA: Louisa Co.,
South Anna River, Rte 657, (csuc); North Anna
River, Rte 601, (csuc); Kansas: Chatauqua Co., Big
Caney River, 1 1/2 mi W of Elgin, (csuc); Texas:
Caldwell Co, Plum Creek at US Hwy 90 (rwe);
Goliad Co., 3 mi N of Goliad, (Rwa); Gonzales Co.,
Guadalupe River, 4 mi S of Gonzales (RWG);
Palmetto State Park, 6 mi S of Luling, nr. US Hwy
183, (RWG); Hidalgo Co., Bentsen, Rio Grande State
Park, (csuc); McLennan Co., Hog Creek, (RWG);
Real Co, Nueces River at Tex Hwy 55, S of
Barksdale, (RwG); Reeves Co., roadside irrigation ditch
W of Balmorhea (RwG); Williamson Co., 3 mi SW of
Taylor, (RWG); Mustang Creek by Carlos G. Parker
Blvd (= Loop 427), Taylor, (RwG); Cororapo: Las
Animas Co., Purgatory River, Bent Cyn nr. Delphi
(csuc, RWG); New Mexico: Guadalupe Co., Santa
Rosa, 4600 ft. (ummz); ARIZONA: Maricopa Co., Salt
River at Corn Bluff Campground, 15 mi NE Mesa, (-
FSCA); Mexico: Nuevo Leon: in foothills of the north-
ernmost range of mountains, (UMMZ); Apodaca, just
NE of Monterrey, near course on Instituto farm, (-
UMMZ, RWG); Chihuahua: La Cruz, (RWG); Naica
(ummz); Durango: 3.7 mi NW of Gomez Palacio on
Hwy 49, 3700 ft, (ummz); 8.2 mi N of Gomez
Palacio on Hwy 49, (ummz); Rio Nazas at Mex. Hwy
49, 3.4 mi S Ciudad Leon Guzman, 4200 ft. (pre).
Erpetogomphus sipedon Calvert
(figs. 19-thx, 39, 40-abd, 61-hamules, 83-penis,
105, 106-app, 158, 159-vertex, 168-leg, 187, 188-
vl, 203-distr)
Erpetogomphus sipedon Calvert, 1905: 165 (descr. 2 Jalisco:
Guadalajara). — Calvert 1907: 399 (9 ‘Las Bocas in
Durango...’); Calvert 1908c: xxx (Erpetogomphus sipedon,
type 2, Guadalajara, found by Schumann); Calvert
1909: 481 (Cuernavaca, Mexico); Williamson and
Williamson 1930: 13 (summary of status); Kimmins
1969: 297 (type in BMNH); Paulson 1982: 256 (Mex.);
Davies and Tobin 1985: 28 (cat.); Tsuda 1986: 87 (cat.);
Bridges 1991: VII.195 (cat.); Tsuda 1991: 95 (cat.).
GARRISON: Revision of Erpetogomphus
Description
Male. — Entire face light green; this colour more vi-
vid on frons, post- and anteclypeus; base of mandible,
labrum light green, vertex with brown around and on
ocellar prominences, remainder light green; antennal
pedicel dark brown dorsally, light green ventrally; se-
cond segment and flagellum dark brown; occiput
light green, crest slightly emarginate medially, its
hind margin covered with long brown hairs; rear of
head brown, darker toward occipital foramen, beco-
ming light green laterally.
Prothorax predominantly dark brown, becoming
light green dorsally; anterior and posterior lobes light
green. Synthorax entirely light green (probably a viv-
id apple green in life) with following dark brown: an-
tehumeral stripe ending dorsally before antealar sinus,
narrow stripe along humeral suture, this stripe widen-
ing at dorsal 0.30 of suture and with anterior margin
just touching upper part of antehumeral stripe, ante-
rior 0.50 of mesinfraepisternum. Coxae, trochanters
ivory, femora light green with black along external
surfaces, more so on mesofemora; most of lateral sur-
face of mesofemora dark brown, these markings more
restricted and limited to apical 0.30 of metafemora;
tibiae black with small lateral ivory stripes along basal
0.30; tarsi and armature black.
Wings hyaline, venation dark brown, basal wing
venation brown, anterior margin of costa yellow; pte-
rostigma brown, black along margins.
Venational statistics (n = 13). Fifth antenodal
thickened in all wings; number of marginal cells be-
hind fore wing paranal cells: 1; anx: fore wing 12-
13/12-15, hind wing 9-10/9-10; pnx: fore wing 7-
10/7-9, hind wing 9-11/8-10; cs under pterostigma:
fore wing 5-6/5-6, hind wing 5-6/6; anal triangular
cells: 4. Hind wing 28-31 mm.
Abdomen (fig. 39). Segment 1 light green with
dorsolateral brown stripe; segment 2 similar to seg-
ment | but dark brown expanding posteriorly behind
auricle, contracting again and interrupted just before
brown annulus; segment 3 similar to segment 2 but
with dark brown at transverse carina, anterior end of
dorsolateral not on or just touching segment 2, its
posterior end broadly connecting with dark brown
annulus; segments 4-6 similar to segment 3 with pos-
terior 0.40 wider than anterior 0.60; segment 7 large-
ly pale except for brown along transverse carina, be-
coming red brown posteriorly, especially so laterally;
segment 8 largely red brown with ill-defined light yel-
low brown middorsal and ventrolateral stripes; seg-
ment 9 similar to segment 8; segment 10 red brown
basally, becoming pale yellow brown distally.
Abdomen 33-37 mm.
Cercus (figs. 105-106) ivory, strongly angulate
near middle; tip of cercus beyond angulation narrow,
drawn out to a fine point; ventral margin with inferi-
221
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
or carina at basal 0.30. Epiprocts yellow brown, typi-
cal of genus.
Accessory genitalia (fig. 61). Anterior hamule
black, divided at upper 0.30, lower branch forming a
small, truncate appendage; posterior hamule white,
triangular, with anterior basal shoulder, tip with
small, anteriorly recurved tooth; penis with small ser-
rated lateral lobes, membranous hood short, exposing
a subcutaneous membrane which forms posterior
margin of segment (fig. 83); cornuae (fig. 83) moder-
ately long, widely separated, medial lobe present, usu-
ally 0.5 to 1.0 length of outer lobe.
Holotype female. — Face entirely pale except for
small black area at base of mandible, occiput (fig.
159) dark brown around ocelli, remainder pale.
Prothorax mostly pale, darker along pleura.
Synthorax (fig. 19) mostly pale except for brown mar-
gin around antealar sinus and incomplete antehumer-
al stripe, its upper end joined with narrow humeral
stripe. Coxae, trochanters, and femora pale with api-
cal 0.30 black, especially mesofemora, metathoracic
legs with black less extensive (fig. 168), forming black
streaks externally; tibiae entirely black except for pale
line laterally, tarsi and armature black.
Abdomen (fig. 40) similar to male but with dark
brown dorsolateral stripe of uniform width on seg-
ment 2, segments 8-9 mostly black dorsally, sides
tawny, annulus pale, cerci lacking. Abdomen 35 mm.
Venational details: number of marginal cells be-
hind fore wing paranal cells: 2/2; anx: fore wing
13/13, hind wing 9/11; pnx: fore wing 9/9, hind
wing 10/12; cs under pterostigma: fore wing 6/7,
hind wing 7/6. Hind wing 31 mm.
Vulvar lamina (fig. 187) with broadly rectangular
flaps meeting at anterior margin; juncture of Y-
shaped postlamellar ridge at level of posterior margin
of lamina.
Diagnosis
Males of Erpetogomphus sipedon are most similar to
E. heterodon but are easily distinguished by the struc-
ture of the penis and cercus. The cornua in £. sipedon
has a medial lobe (fig. 83), which is lacking in £. he-
terodon (fig. 87), and the tip of the cercus in E. sipe-
don are slender and finely attenuate (figs. 105-106),
not robust and short as in E. heterodon (figs. 110-
111). Erpetogomphus heterodon and E. sipedon are the
only two species of the E. designatus group which sha-
re a similar thoracic design. However, E. sipedon lacks
any dark thoracic stripes on the second and third la-
teral sutures; a vestigial incomplete second lateral stri-
pe and complete narrow third lateral stripe exist on E.
heterodon. The thoracic pattern of E. boa is also simi-
lar to E. sipedon, but E. boa is well differentiated by
genital characters given in the keys. Although the dif-
ference in somatic patterns between £. lampropeltis
DOD
and E. sipedon is great, these two species are separable
by the structure of the penis. In £. sipedon, the hood
is abbreviated, exposing a subcutaneous membrane,
which latter forms the posterior margin of the seg-
ment. In E. lampropeltis, the hood is of normal length
and it forms the posterior margin of the segment.
Although females of E. sipedon are easily differen-
tiated from all other similar species by differences in
. colour and pattern, their separation from the same
species using only structural characters is more diffi-
cult. Species morphologically similar to E. sipedon in-
clude E. crotalinus, E. compositus, E. viperinus, E.
bothrops, and E. heterodon.
The occiput of £. sipedon is straight or, at most,
weakly notched, (notch in E. crotalinus not clearly
seen in the paralectotype female, [fig. 161]). The lon-
gitudinal distance of the occiput (hereafter called
width) is narrower in £. sipedon than in E. crotalinus,
and the posterolateral ocellar tubercle is less pro-
nounced in £. sipedon than in E. crotalinus.
The occipital crest of E. compositus is sinuous
(mostly linear in Æ. sipedon) and the postocciput is
easily visible in dorsal view (fig. 163); the same struc-
ture is barely visible in E. sipedon (fig. 159).
The structure of the postlamellar ridge provides the
best means of separation between E. sipedon and sister
taxa E. bothrops and E. viperinus. In E. sipedon, this Y-
shaped structure is short: the juncture of the lateral
arms meets at or near the hind margin of each vulvar
plate. The longitudinal stem also does not possess any
circular or oval depressions on either side (figs. 187-
188). In E. bothrops and E. viperinus, the juncture of
the Y-stem is well posterior to the hind margin of
each vulvar plate, and a well-defined oval or circular
depression exists on either side of the longitudinal
ridge (figs. 183-184).
I have been unable to find any morphological dif-
ferences between females of E. sipedon and E. hetero-
don. Body pattern (figs. 19, 23) and distribution (figs.
201-204) will serve to separate females of these close-
ly related species.
Remarks
Variation. — The male description is based on 11
males from Jalisco, Morelos, and Durango states,
Mexico. The four from Durango are slightly larger
(hind wing 30-31 mm); and their thoracic pattern is
similar to that of the holotype female (fig. 19). The
other seven males are smaller (hind wing 28-29 mm),
and the lower 0.60 of the dark humeral stripe is nar-
rower. The Durango males, though larger, have a
sparser venation. For example, three have no margin-
al cells behind fore wing paranal cells, one has one
marginal cell in both wings; and all other specimens
have one marginal cell, except for the left fore wing of
one Acatlipa male, which has no marginal cells. Two
of the Durango males have a 3-celled anal triangle in
both hind wings, while all others possess 4 cells. The
cerci of the Durango males are less angulate and the
tips more robust (fig. 106) than the other males, but
other characters of the body and penis leave no doubt
that they represent £. sipedon.
Variability for females is less pronounced than for
males, and I can find no overall differences in vena-
tional patterns among the 12 females examined.
Female venational statistics (n = 12, including hol-
otype): number of marginal cells behind fore wing
paranal cells: 1-3/0-3; anx: fore wing 12-14/12-15,
hind wing 9-12/9-11; pnx: fore wing 8-10/8-10, hind
wing 9-11/8-12; cs under pterostigma: fore wing 5-
715-7, hind wing 5-7/4-6. Hind wing 30-34 mm.
Abdomen 36-37 mm.
When Calvert (1905) described this species, he had
only 6 females available; and he noted differences in
the condition of the occiput between the holotype
and a paratype from Matamoros, Puebla (figs. 158-
159). Both forms were represented in his key. I can-
not consider the differences noted by Calvert as use-
ful, as much variation seems to exist in the shape of
the occipital ridge in females of this and other species.
At the time of the original description, Calvert
(1905) postulated that the female of E. sipedon could
be E. boa, which was then known only from one in-
complete teneral male. The discovery of more speci-
mens of both species, and especially the capture of
both males and females of Æ. sipedon in Durango,
leaves no doubt that both are valid species.
Biology. — González (in litt.) collected this species
at Palo Bolero, where adults perched on herbaceous
vegetation on the banks of the Rio Sabinos. Its
known distribution suggests that it is more widely
distributed, but specimens are rare in collections.
Specific ecological requirements may restrict its oc-
currence to certain sites, but a more likely answer to
its apparent rarity is a lack of thorough collecting
within its range. Collection dates range from 21 May
(Cuernavaca) to 13 August (Nombre de Dios).
Distribution (fig. 203). — Erpetogomphus sipedon is
known from northern Durango (Las Bocas [Calvert,
1907] settlement on the Rio Florida in extreme
northern Durango near Villa Ocampo, approx.
26°25°N 105°25’W, [Selander and Vaurie 1962])
south through Morelos and Puebla (approx.
18°50’N, 99°W) and seems restricted to the highland
area of central and western Mexico from altitudes of
1000 m (Palo Bolero) to about 2073 m (Las Bocas).
González (in litt.) collected this species in Mexico
state (Tonatico) during July, 1992.
Material
Type data. — Holotype female: white printed label:
‘[Mexico] Guadalajara,/Jalisco,/July. Schumann’,
GARRISON: Revision of Erpetogomphus
white, partially printed label: “Erpetogomphus 2 /sipe-
don Calv. Tyre /P. P. Calvert, det. 1905/B. C. A.
Neur., p. xxx, 166/orig. of Pl. VIII, ff. 34, 40° with
‘(form/a)’ handwritten on left side; printed label
‘Brit. Mus./1911-339.’; small round label with red
margin: “Holo-/type’.
Other material (118, 122, including holotype
?).- Mexico: Durango: Nombre de Dios, just SE of
Durango on Mex. Hwy 45 (23°51’N, 104°14’W),
5900 ft. (1800 m), 13 Aug. 1947 (W. Gertsch, M.
Cazier), 46,79 (AMNH, RWG); Morelos: Cuernava-
ca, 21 May 1898 (no collector), 19 (curc); Cuerna-
vaca, 7 July 1900 (C.C. Deam), 29 (ummz); Palo
Bolero, km. 18.3, route 95, 5 km. S. Acatlipa, 1000
m (approx. 18°45’N, 99°15’W), 26 June 1985 (E.
Gonzalez), 26 (UNAM); 18 June 1986 (E. Gonzalez),
26 (UNAM), 12 July 1986 (E. González, V. Garcia),
3d (rwe); Morelos: Pueblo Cocyotla, Hwy 421,
about 30 km SW of Cuernavaca, 23 July 1992 (J.
Daigle), 16 (jp); Puebla: Matamoros, no date, (Otis
W. Barrett), 19 (ANSP).
Erpetogomphus lampropeltis Kennedy
Description
Male. — Face entirely pale with dark markings as
follows: along basal margin of labium, along fronto-
clypeal suture (less developed in £. / natrix), base of
antefrons; vertex all dark brown; antennae dark
brown; anterior margin of vertex with an arcuate pit,
one each anterolaterad to median ocellus; area be-
tween these pits tumid (fig. 138), occiput white
green, wide, slightly tumid medially, crest entirely
black, slightly raised, linear or slightly convex posteri-
orly, some with a small median notch, its hind mar-
gin covered with pale brown hairs; postocciput slight-
ly concave, pale; rear of head dark brown.
Prothorax largely brown with pale green or grey
green areas on anterior and posterior lobes, and with
two small median spots on median lobe; contrasting
dark/pale areas more prominent in nominotypic sub-
species; pale colours of synthorax grey or grey green
(E. 1. lampropeltis) to green (El. natrix) with follow-
ing dark brown areas: middorsal stripe, widening ba-
sally to collar, its upper end covering antealar sinus,
extending posteriorly and connecting with broad,
well defined antehumeral and humeral stripes; well
defined sinuate second lateral stripe; third lateral
stripe. Coxae, trochanters pale grey green with slight
wash of brown posteriorly; tibiae pale grey green with
defined superior surfaces brown, becoming dark
brown distally; tibiae black with narrow strip of pale
grey green along basal 0.30 to 0.50 of lateral margin;
base and armature black.
Wings hyaline, basal wing venation brown; anteri-
or margin of costa yellow up to pterostigma; pteros-
223
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
tigma black; fifth antenodal thickened in all wings
(with rare exceptions).
Abdomen. Abdominal segment 1 pale grey green
to green with basal 0.25 brown and with dorsolateral
brown stripe; segment 2 with dorsolateral stripe send-
ing a ventral stem posterior to auricle and (often) an
isolated spot on anteroventral margin of segment,
dorsolateral stripe often connecting with black annu-
lus or ending just before; segment 3 with isolated
black dorsolateral spot surrounding lateral carina; its
anterior end tapering and sometimes touching anteri-
or margin of segment, distal 0.25 of segment with
dorsolateral black increasing in width posteriorly and
touching dorsally near black annulus; segment 4 sim-
ilar to segment 3, but two black spots broadly con-
nected forming a continuous dorsolateral stripe occu-
pying distal 0.80 of segment, this stripe often
constricted in middle; segments 5 and 6 similar to
segment 4, but with middorsal pale areas becoming
darker; segment 7 with anterior 0.50 pale, except for
black transverse carina, becoming entirely tawny to
black posteriorly; segments 8-10 yellow brown, be-
coming black dorsally in most specimens; foliar ex-
tensions dark brown to black.
Cercus (figs. 107-108) ocher, in lateral view slight-
ly concave dorsally near base, posterior 0.25 moder-
ately angulate, the short tips slightly recumbent; cer-
cus enlarged ventrally at basal 0.40 to 0.50 with
inferior carina along same area, remainder slightly
concave with tip forming a blunt point. Epiprocts
yellow brown, gently curved dorsally at 90°, tips (fig.
132) dorsoventrally flattened and divergently bluntly
pointed.
Accessory genitalia (figs. 62-63). Anterior hamule
dark brown, divided at basal 0.50, lower branch
forming a small, posteriorly directed truncate ap-
pendage; posterior hamule grey white, triangular,
with anterior basal shoulder, tip bluntly rounded and
armed with a small anteriorly directed tooth on pos-
terior border; penis with lateral lobe small, roughly
semicircular, serrate; cornuae (figs. 84-85) moderate-
ly widely separated, tips broadly rounded with well
developed median lobes.
Female. — Head as in male, but with crest of occi-
put slightly more erect and convexly arcuate; posteri-
or margin of postocciput slightly concave; prothorax
and thorax as in male, but with pale areas generally
more extensive; wings hyaline or with saffron infu-
sion at base in some E£.l. natrix, abdomen in well pre-
served specimens with pale colours primarily white
with dull orange dorsally; black dorsolateral stripes
and spots more reduced than in male, these stripes
sometimes separated or greatly constricted on seg-
ments 4 and 5; cercus ocher.
Vulvar lamina (figs. 189-190) with prominent ba-
sal plates, quadrangular laminar plates acuminate
224
posteriorly, meeting at anteromedial margin; cleft
between plates forming a 90° angle; V juncture of Y-
shaped postlamellar ridge at level of posterior margin
of lamina.
Diagnosis
This species is most similar to its congeneric relati-
ves, E. sipedon and E. compositus, and is diagnosed un-
_ der those species.
Remarks
The original description of £. lampropeltis is brief
and is based on an unspecified number of males and
females from Sespe Creek, Fillmore, Ventura
County, California. Williamson and Williamson
(1930) originally described £. natrix in detail from 21
males and two females from Baja California, Mexico.
An abstract of their diagnosis of males £. natrix from
E. lampropeltis is as follows:
‘The head and thorax of /ampropeltis are duller and paler
than in matrix... the rear of the head is brown in /ampropel-
tis and black in matrix; the thoracic brown is paler in lampro-
peltis and the pale areas are grayer, giving much less contrast
in the thoracic pattern; the two lateral dark stripes are always
joined, often very broadly in lampropeltis, and are not joined
in.....natrix,....the femora are brown in /ampropeltis.....black
in natrix, and the inferior dilated edge of abdominal seg-
ments 8 and 9 is black in lampropeltis and, in natrix, is col-
ored light vivid brown like the sides of the segment adjacent
to the edge.....’
The apparent differences in morphology described
above fall well within the latitude of individual and
geographic variation and do not, in my opinion, con-
stitute specific differences. Further, several specimens
of E. natrix from various parts of its range have mac-
ulation patterns more closely approaching those of £.
lampropeltis. For example, the antehumeral and hu-
meral stripes are briefly connected, the dorsum of ab-
dominal segments 7-10 have black as in E. lampropel-
tis, and the inferior dilated margin of abdominal
segments 8 and 9 is black. I can also detect no differ-
ences in the shape of the vulvar lamina between fe-
males of E. lampropeltis and E. natrix.
The forms are allopatric, separated by the
Colorado Desert. The large series of E. lampropeltis
(mostly topotypes) that I examined does show consis-
tent differences in pale thoracic colouration and mi-
nor differences in thoracic maculation. I consider the
forms to represent subspecies separable as follows:
1. Pale thoracic colouration grey green, antehumer-
al and humeral stripes broadly connected on dor-
sal 0.25, after isolating a small pale spot below an-
tealar sinus; second lateral stripe about twice as
wide on dorsal 0.5 and often connecting with
narrow third lateral stripe; dark thoracic stripes,
especially second and third, often not well de-
fined (fig. 20); Ventura, San Diego, Los Angeles,
and western San Bernardino counties, California
E. lampropeltis lampropeltis
— Pale thoracic colouration vivid green, antehumer-
al and humeral stripes not connected, or only
narrowly so; second lateral stripe narrower than
in E. /. lampropeltis, not connecting with third
lateral stripe (fig. 21); central Baja California,
Arizona, New Mexico, western Texas, south
tO NID TANO MEXICO. AA ES
Erpetogomphus lampropeltis lampropeltis Kennedy
(figs. 20-thx, 62-hamules, 84-penis, 107-app, 132-
ept, 138-base of postfrons, 160-vertex, 189-vl, 204-
distr)
Erpetogomphus lampropeltis Kennedy, 1918: 297 (descr. 3
Sespe Creek, Fillmore, Ventura Co., California). —
Needham and Heywood 1929: 80 (descr. d, ©);
Williamson and Williamson 1930: 13 (summary of stat-
us); Needham and Westfall 1955: 147 (descr.); Pritchard
and Smith 1956: 116 (key); Musser 1962: 14 (notes on
larva); Borror 1963: 104 (common name); Montgomery
1968: 133 (distr.); Paulson and Garrison 1977: 157
(Calif.); Paulson 1982: 255 (U.S.); Davies and Tobin
1985: 27 (cat.); Tsuda 1986: 87 (cat.); Bridges 1991:
VII.115 (cat.).
Herpetogomphus lampropeltis. — Byers 1928: 5 (larva un-
known).
Erpetogomphus lampropeltis lampropeltis. — Tsuda 1991: 95
(cat.).
Description
Male. — Colouration and maculation as given un-
der species account, key and in fig. 20.
Venational statistics: number of marginal cells be-
hind fore wing paranal cells: 0-3/1-3. — anx: fore wing
9-14/10-14, hind wing 8-10/8-10; pnx: fore wing 7-
9/5-10, hind wing 7-12/7-12; cs under pterostigma:
fore wing 4-5/4-5, hind wing 3-5/3-6; number of
anal triangular cells: 3-4/3-5. Hind wing 28-31 mm.
Abdomen 34-37 mm.
Female. — Venational statistics (n = 6): number of
marginal cells behind fore wing paranal cells: 0-4/1-3;
anx: fore wing 13-15/12-16, hind wing 9-11/9-11;
pnx: fore wing 9-11/9-10, hind wing 9-12/9-12; cs
under pterostigma: fore wing 5-6/5-6, hind wing 5-
6/5-6. Hind wing 32-36 mm. Abdomen 33-36 mm.
Diagnosis
See account under £. compositus and key to subspe-
cies of E. lampropeltis.
Remarks
Variation. — Venational details of holotype male:
number of marginal cells behind fore wing paranal
GARRISON: Revision of Erpetogomphus
cells: 1/2; anx: fore wing 12/12, hind wing 9/9; pnx:
fore wing 8/9, hind wing 9/9; cs under pterostigma:
fore wing 5/5, hind wing 5/4; number of anal trian-
gular cells: 4/5. Hind wing 28 mm.
The large series of specimens primarily from the
type locality shows little somatic colour and/or pat-
tern variation, due, possibly, to its limited distribu-
tion. The fifth antenodal in the fore wings is general-
ly thickened, but 7 of 40 wings of 20 males examined
have the fourth antenodal thickened; and one wing
has the sixth antenodal thickened. A 3-celled anal tri-
angle is common in this subspecies, as almost half
(19/40) of the wings possess this condition.
The female from Matillija Hot Springs has flaves-
cent wing bases.
Biology. — At the type locality, this species was
abundant during early September, 1982. I collected
54 males within about two hours. All preferred to sit
on exposed rocks and were easily collected with a net.
Hundreds of cast skins were attached to the exposed
rocks. I did not find the species in smaller rivulets
emptying into Sespe Creek. The only female I saw
and collected was found dead in the water.
I returned to the locality in September, 1984, but
the entire area had been fenced off. Five males were
collected in the general vicinity, below Matillija Hot
Springs, but they were rare and more warty.
This is a late summer species: the specimens from
San Bernardino Co. collected in June are all teneral.
Flight dates for specimens from Ventura Co. range
from 10 July to 8 September.
Distribution (fig. 204). — The nominotypic sub-
species has been collected only at three localities in
Ventura County one in Sar Bernardino County, one
in Los Angeles County, and one in San Diego
County, California. It has the most restricted distri-
bution of any Odonata form in southern California.
Material
Type data. — Holotype male: caLIF: Ventura Co.,
Sespe Greek, Eillmores At MOIS RCE
Kennedy); in usnm. I have illustrated the thorax (fig.
20), caudal appendages (fig. 107), and penis (fig. 84)
of the holotype.
Other material (704, 69, including holotype ó).
— U. S. A.: CALIFORNIA: Los Angeles Co., San Gabriel
Mtns., jet of N and W Fork of San Gabriel River, E
of Hwy 39, 11 Aug. 1986 (D. Swinney), 1d (rwe),
Ventura Co., Sespe Creek, 4.0 mi N of Fillmore, 600
ft., 8 Sept. 1971 (D. Paulson), 2d (cc, rwe); 5 Sept.
1982 (R. W. and J. A. Garrison), 546, 19 (rwG);
Matillija Hot Springs, 14 Aug. 1927 (T. Craig), 16,
1? (rsca); just below Matillija Hot Springs, 1400 ft.,
2 Sept. 1984 (R. W. and J. A. Garrison), 56 (Rwa);
Wheeler Hot Springs, 10 July 1927 (T. Craig), 4d,
(rsca); San Bernardino Co, Deep Creek Public
225
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Camp, 15 June 1957 (A. Menke, L. Strange), 1d,
49
(Lacm); San Diego Co., La Jolla, La Jolla Indian Res.
Campground, San Luis Rey River, 25 Aug. 1991 (W.
F. Mauffray), 16 (rwe).
Erpetogomphus lampropeltis natrix Williamson and
Williamson stat. n.
(figs. 21-thx, 63-hamules, 85-penis, 108-app, 1990-
vl, 204-distr)
Erpetogomphus natrix Williamson & Williamson,1930: 19 (-
descr. Sd, 9 Baja Calif. Sur, Purissima, 12 October
1923). — Byers 1939: 50 (quotes Williamson &
Williamson, 1930); Fraser 1940: PI. 6 (penis); Needham
& Westfall 1955: 148 (descr. 6); Borror 1963: 104
(common name); Montgomery 1968: 133 (distr.);
Paulson 1982: 255 (U.S., Mex.); Davies and Tobin 1985:
28 (cat.); Tsuda 1986: 87 (cat.).
Erpetogomphus lampropeltis. — Tinkham 1934: 215 (Tex.),
Needham & Westfall 1955: 147 (Tex.); Gloyd 1958: 8
(Tex.); Kormondy 1960: 122 (Ariz.); Gonzalez & Novelo
1991: 97 (Mex.); Novelo & Gonzalez 1991: 154 (descr.
larva, Mex.).
Erpetogomphus lampropeltis natrix. — Bridges 1991: VII.144
(cat.); Tsuda 1991: 95 (cat.). [Demotion of E. natrix to
subspecies status was advised by me in litt. to Bridges and
Tsuda.]
Description
Male. — Colouration and maculation as given un-
der species account, key, and fig. 21.
Venational statistics: number of marginal cells be-
hind fore wing paranal cells: 0-3/0-2; anx: fore wing
11-15/11-14, hind wing 9-11/9-11; pnx: fore wing 6-
11/7-10, hind wing 8-14/7-12; cs under pterostigma:
fore wing 4-6/4-6, hind wing 4-6/4-6; number of
anal triangular cells: 3-4/2-4. Hind wing 24-33 mm.
Abdomen 30-37 mm.
Female. — Venational statistics (n = 17, including
allotype): number of marginal cells behind fore wing
paranal cells: 0-6/0-5; anx: fore wing 12-15/12-15,
hind wing 9-11/9-11; pnx: fore wing 7-11/7-12, hind
wing 9-12/9-12; cs under pterostigma: fore wing 4-
6/4-7, hind wing 4-7/5-7. Hind wing 30-35 mm.
Abdomen 36-41 mm.
Diagnosis
See comments under E. compositus and key to sub-
species of E. lampropeltis.
Remarks
Venational details of holotype male: number of
marginal cells behind fore wing paranal cells: 2/1;
anx: fore wing 13/13, hind wing 9/9; pnx: fore wing
9/9, hind wing 10/9; cs under pterostigma: fore wing
4/4, hind wing 5/4; number of anal triangular cells:
4/3. Hind wing 30 mm.
226
Venational details of allotype female: number of
marginal cells behind fore wing paranal cells: 2/1;
anx: fore wing 13/13, hind wing 9/10; pnx: fore wing
7/8, hind wing 10/9; cs under pterostigma: fore wing
6/5, hind wing 5/5. Hind wing 30 mm.
Variation. — Intrapopulational variation of body
maculation appears to be as great as interpopulation-
al variation. The antehumeral and humeral stripes
_ may be connected or separate in any small series from
a given locality: 1 of 3 specimens from Cane Springs,
Mojave Co., 3 of 12 specimens from Cave Creek,
Maricopa Co. (4 have these stripes barely connected),
and 2 of 11 specimens from 15 mi E of Douglas,
Cochise Co., Arizona, all have connected antehumer-
al and humeral stripes. Similar degrees of variation ex-
ist for the dorsal and dorsolateral darkening of ab-
dominal segments 7-10. Some specimens have little
black on these segments (as noted by Williamson and
Williamson, 1930, for their type series of E. natrix),
but others have various degrees of darkening, and
some specimens are as dark as I have observed for no-
minotypic E. lampropeltis.
A male I collected at Las Parras, Baja California,
has the posterior 0.5 of the dorsum of abdominal seg-
ments 7 and all of segments 8-9 with black. The male
from ‘Corralitos’ is unusual in having narrow second
and third lateral thoracic stripes. The upper 0.5 of the
second lateral stripe is also lacking, but in all other as-
pects, this specimen is E. lampropeltis natrix by mor-
phology and overall body colouration and macula-
tion.
Biology. — Williamson and Williamson (1930) de-
scribed the type locality. González (in litt.) collected
this subspecies with E. crotalinus in Durango state (La
Michiliä). I collected one male of the species at Las
Parras in September, 1985 (Garrison, 1986) as it sat
upon an exposed rock in the middle of the small
stream. It was the only individual I saw.
This subspecies can be common at certain sites in
Arizona and New Mexico. Jo A. Garrison and I col-
lected E. lampropeltis natrix on the East Fork of the
Gila River at Grapevine Campground, where it was
equally as common as E. heterodon. Adults landed on
exposed rocks, exposed bleached logs in the center of
the stream, or on the gravel shores. I have also taken
it on sandy-bottomed desert streams, often in compa-
ny with Æ. compositus. Novelo (pers. comm.) found
this subspecies with the more common E. crotalinus
in Durango state (La Michiliá).
I observed oviposition on 1 August 1992 at the
Agua Fria River, Yavapai County, Arizona. Females
appeared over the stream surface in the late afternoon
(about 6: 15 p.m.), and flew rapidly over the water,
dispersing their eggs by dipping the abdomen briefly
into the water. This behaviour continued until sun-
down (about 7: 30 p.m.). A few males were still
present, sitting on emergent rocks near oviposition
sites. No copulations were observed.
Altitudinal gradients for the species range from
near 923 m (Tangle Creek, Yavapai Co., Ariz.) to
1754 m in New Mexico and southeastern Arizona (W
of Portal). Like E. / lampropeltis, E. |. natrix is pri-
marily a late summer to autumn species. Flight dates
range from 27 June to 10 October.
Distribution (fig. 204). — Though originally de-
scribed from Baja California, £. lampropeltis natrix is
widely distributed throughout the southwestern
United States. It has been collected as far east as west-
ern Texas (Tinkham, 1934, Gloyd, 1958) and
Durango and Chihuahua, Mexico. The most north-
erly records are from northcentral Arizona. I have
seen no specimens from central or eastern New
Mexico, although it probably occurs in those areas.
The species apparently does not penetrate north of
the Mogollon Rim in Arizona northwest into the
Colorado Desert.
Material
Type data. — Holotype male: Mexico: Baja California
Sur: Purissima, 12 Oct. 1923 (JJ. H. Williamson); allotype
female, same data but 6 Oct. 1923 (both examined), in
UMMZ. I have illustrated the thorax (fig. 21), hamules (fig.
63), caudal appendages (fig. 108), and penis (fig. 85) of the
holotype, and vulvar lamina (fig. 190) of the allotype.
Other material (1016, 229, including holotype d and
allotype 2). — U. S. A.: ARIZONA: Cochise Co.: Bear Creek,
Rte 61, 6 mi SE Parker Lake, Coronado National Forest
(csuc); mouth Miller Canyon, Huachuca Mtns., (Fsca,
UMMZ), pond N of Slaughter Ranch, 15 mi E of Douglas,
3800 ft, (RWG); springs just E of Slaughter Ranch, 17 mi E
of Douglas, (RWG); Bear Creek, Rte 61, 6 mi SE Parker
Lake, Coronado Nat’! Forest, (csuc), San Pedro River at
Hwy 90 (rsca); San Pedro River, jct. Charleston Rd.,
(Byuc); San Pedro River, 9 mi SW of Tombstone (Byuc);
San Pedro River at Hereford Rd., (RwG); Southwestern
Research Station, 5 mi W of Portal at John Hands Picnic
Grounds, 5400 ft, (rsca); Gila Co., Tonto Natural Bridge
(Rw6); Webber Creek at Camp Geronimo, 8 mi ENE of
Payson, (RwG); Maricopa Co., Cave Creek by Ocotillo Rd,
Cave Creek, (Rwa); Camp Creek by Cave Creek Rd., 12 mi
NE of Scottsdale Rd., 2700 ft., Tonto Nat'l. Forest, (RWG);
Sycamore Cyn., 3.3 mi WNW of Hwy 87 (rwe); Mojave
Co., irrigation ditches E of Cave Springs, (DRP); Pima Co.,
Sabino Cyn, N of Tucson (rsca); Lower Sabino Cyn,
(RwG); Upper Sabino Cyn, ca. 3 mi N of visitor center, N of
Tucson (rwG); Santa Cruz Co., Sonoita Creek Ranch (on
way to Salero Cyn), along Santa Cruz River, along Ariz.
Hwy 82, 2 mi SW of Patagonia, (RwG); White Rock
Campground, Peña Blanca Lake, 4200 ft, (rwe, JD);
Yavapai Co., Agua Fria River at Black Canyon City, (RWG);
Tangle Creek, nr Tangle Creek cabin, ca 33 mi N of
Carefree, TIN-RSE, sec. 1, 2800 ft (Rwa); on limestone rim
of Montezuma’s (sic) Well (National Monument) (RwG);
Sycamore Canyon, Atasco Mtns (LACM); New Mexico:
Catron Co, Gila Cliff Dwellings, (rwa); West Fork Gila
River at Gila Cliff Dwellings Natl Mon., 5700 ft, (RWG);
Grant Co., Gila River at Cliff (rsca); Mangus Springs Creek,
GARRISON: Revision of Erpetogomphus
Hwy 180, Mangus Springs, (RWG); Grapevine
Campground, East Fork of Gila River at N. Mex. Hwy 15,
5600 ft, (Rwc); Texas: Jeff Davis Co., Ft. Davis, (rsca);
Limpia Creek, 2-3 mi N of Ft. Davis, (ummz); Musquiz
Creek, 6 1/2 mi SE Ft. Davis (rsca); Mexico: Chihuahua:
Cascada de Basaseachic National Park, (csuc, Byuc); San
Diego Canyon, (Byuc); Durango: Arroyo El Temazcal (ca-
mino a El Aleman), Reserva de la Biosfera La Michiliá,
(UNAM); km 21 Camino Suchil-Reserva de la Biosfera La
Michiliá, ‘Corralitos’, (UNAM); Sonora: Maycoba River,
(Byuc); Maycoba River, W of Maycoba, (Byuc); 8 mi W of
Maycoba River, (csuc); Maycoba River, Hwy 16, 13 mi W
of Maycoba at San Francisco Bridge, (csuc); Rio
Magdalena, Hwy 15 S of Immuris, (BYUC, csuc); small riv-
er W of Vecora, Rte 15, (csuc, Byuc); Baja California Sur:
Purissima (Fsca); Las Parras, about 18 km W of Hwy 1 on
route to San Javier, SW of Loreto, (RWG).
Erpetogomphus crotalinus (Hagen in Selys)
(figs. 22-thx, 41-abd, 64-hamules, 86-penis, 109-
app, 133-ept, 161-vertex, 169-leg, 191, 192-vl, 205-
distr)
Ophiogomphus crotalinus Hagen in Selys, 1854: 40 (21 sep.)
(3, 2 descr. from Mexico).
Erpetogomphus crotalinus. — Hagen in Selys 1858: 332 (72.
sep.) (descr. d, 9, relationship with E. menetriesii);
Walsh 1863: 253 (mentioned); Selys 1869: 174 (11 sep.)
(Mex., attributes authorship to Selys); Selys 1873b: 519
(75 sep.) (list, attributes authorship to Selys); Kirby 1890:
61 (cat., attributes authorship to Selys); Calvert 1899: 38
(Tepic, Mex.); Calvert 1905: 165 (Mex.); Calvert 1907:
399 (Mex.), Calvert 1909: 481 (distr. in Mexico);
Muttkowski 1910: 86 (cat.); Ris 1917: 153 (comparison
with £. boa); Kennedy 1918: 298 (comparison with E.
lampropeltis); Byers 1939: 50 (quotes Calvert 1907);
Williamson and Williamson 1930: 12 (summary of sta-
tus); Montgomery 1968: 133 (distr.); Paulson 1982: 255
(cat.); Davies and Tobin 1985: 27 (cat.); Tsuda 1986: 87
(cat., attributes authorship to Selys); Bridges 1991:
VILS4 (cat.); Tsuda 1991: 95 (cat); Cannings and
Garrison 1991: 478 (Mex.); Gonzalez and Novelo 1991:
97 (Mex.); Novelo and Gonzalez 1991: 150 (descr. larva,
Mex.).
Herpetogomphus crotalinus. — Hagen 1875a: 43 (cat.); Selys
1879: 64 (2 sep.) (characters of genus); Kirby 1890: 61
(cat.).
Description
Male. — Labium grey white, base of mandibles pale
green becoming red brown distally, lips black, re-
mainder of face pale green except for narrow wash of
brown at base of antefrons, vertex mostly dark brown
with green medially behind postoccipital tubercles;
antennae dark brown; anterior margin of vertex with
two lanceolate pits, each anterolateral to median ocel-
lus; occiput bright green, broad, gently tumid medial-
ly, crest black, linear to emarginate medially, its rim
covered with row of long brown hairs; postocciput gr-
een, smoothly concave or somewhat angulate medial-
ly, rear of head light green with brown near occipital
foramen.
227
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Prothorax predominantly green, most of middle
lobe and posterior of frontal lobe marked with dark
brown, a small middorsal green spot longitudinally
divided by narrow dark brown line medially on mid-
dle lobe. Synthorax (fig. 22) entirely green with
brown along margins of antealar crest in a few speci-
mens with a hint of a dark brown antehumeral stripe.
Coxae, trochanters pale green, femora green with
streak of dark brown becoming wider distally, these
streaks restricted to lateral surface of femora (fig. 169)
and brown reduced to apical 0.50 of metafemora; a
supplementary narrow, linear streak of dark brown
more posteriorly placed on femora and, in some spec-
imens, touching lateral streaks of brown distally; tib-
iae ranging from all yellow green except dark brown
laterally, to all brown with yellow on lateral carinae;
tarsi and armature black or with some yellow on exte-
rior surfaces of second and third tarsomeres.
Wings hyaline, venation dark brown, anterior mar-
gin of costa yellow to base of pterostigma, this struc-
ture brown, veins bordering it black.
Venational statistics. Fifth, rarely fourth or sixth,
antenodal thickened in all wings; marginal cells be-
hind fore wing paranal cells: 1-3/1-3; anx: fore wing
11-13/11-13, hind wing 8-9/8-10; pnx: fore wing 7-
10/7-10, hind wing 8-10/8-10; cs under pterostigma:
fore wing 4-6/4-7, hind wing 4-7/4-6; anal triangular
cells: 3-4/3-5. Hind wing 26-31 mm.
Abdomen (fig. 22). Segment 1 green, slightly fus-
cous basally, with small line of black at posterior artic-
ulation point; segment 2 all green with narrow dorso-
lateral streak of black above auricle and with poorly
defined dorsolateral brown spot on posterior 0.25 of
segment, dorsal 0.75 to 0.50 of annulus black; seg-
ment 3 pale green, becoming white ventrally with
black along transverse carina and elongate black dor-
solateral spot on posterior 0.25 of segment connect-
ing to black annulus; segments 4-7 similar to segment
3, but with narrow anterior offshoot of black laterally
on transverse carina and with posterior midlateral
black occupying posterior 0.50 of segments, middor-
sal pale area of these segments becoming progressive-
ly more tawny posteriorly, especially so on posterior
0.50 of segment 7; segments 8-9 ocher yellow with
well defined dark brown dorsolateral stripe; segment
10 ocher becoming red brown at basal 0.50.
Abdomen 31-36 mm.
Cercus (fig. 109) yellow, strongly to moderately
angulate dorsally at distal 0.50-0.30; tip of cercus be-
yond angulation attenuate, drawn out to a blunt
point, ventral margin with inferior carina at basal
0.40 to 0.50. Epiprocts yellow brown, gently curved
dorsally at 90°, extending to distal 0.25 or almost to
level of tip of cercus, tips forming divergent points,
dorsoventrally flattened (fig. 133).
Accessory genitalia (fig. 64). Anterior hamule
228
black, divided at upper 0.50, lower branch forming
small posteriorly directed truncate appendage; poste-
rior hamule pale green, triangular, with anterior basal
shoulder, tip rounded with a small anteriorly directed
tooth on posterior margin; penis with semicircular
spinose lateral lobe, cornuae separated; with medial
lobe, and a long narrow flap at outer distal margin,
membranous hood present but not overlapping.
Abdomen 31-37 mm.
Female. — Head as in male but with crest of occiput
(fig. 161) more emarginate medially, in some speci-
. mens with lateral margin arcuate; prothorax and syn-
thorax as in male; legs with black areas more reduced
than in male; wings with slight wash of yellow at base
(more so in tenerals); abdomen as in male but with
black areas more reduced on abdominal segments 1-
3; dorsolateral brown stripes almost complete on seg-
ments 4-7 except for basal 0.05 of each segment,
these stripes complete on segments 8-9; cercus ocher.
Abdomen 30-36 mm.
Vulvar lamina (figs. 191-192) with broadly rectan-
gular flaps meeting at anterior margin, juncture of Y-
shaped postlamellar ridge at level of posterior margin
of lamina.
Venational statistics: number of marginal cells be-
hind fore wing paranal cells: 1-4/1-3; anx: fore wing
11-13/11-13, hind wing 8-10/8-10; pnx: fore wing 7-
10/7-9, hind wing 8-10/8-10; cs under pt: fore wing
4-7/4-6, hind wing 4-6/4-6. Hind wing 26-32 mm.
Diagnosis
Erpetogomphus crotalinus is unique in having the
extensor surfaces of the tibiae yellow (fig. 169). Some
specimens have largely brown tibiae, but the lateral
costate ventral margins are always yellow; all other
congeners have dark brown to black tibiae. The al-
most immaculate synthorax is shared by only two ot-
her species: E. cophias (fig. 27) and some E. elaps (fig.
13), but the male cerci of these two species (figs. 100,
116) are never angulate as in £. crotalinus (fig. 109).
The cornua of the penis (fig. 186) has a well develo-
ped median lobe and a long, acuminate tip, a condi-
tion unique to this species. This species is superficial-
ly similar to E. heterodon and is diagnosed under that
species.
Females of £. crotalinus differ from other species by
three characters of the head, the presence of anterolat-
eral pits at the base of the frons, the relatively broad
occiput, and the medial notch of the crest. The vulvar
lamina (figs. 191-192) provides no discriminatory
characters, and is similar to those of E. lampropeltis
(figs. 189-190), E. compositus (fig. 194), E. heterodon
(fig. 193), and E. sipedon (figs. 187-188).
Remarks
The description is based on 91 males and 43 fema-
les. The lectotype male is reasonably well preserved,
although it lacks the accessory genitalia. I illustrate
the cercus (fig. 109) and right hind leg (fig. 169) of
the lectotype, and the vertex, occiput (fig. 161), and
vulvar lamina (fig. 191) of the paralectotype.
Venational details for lectotype male: number of
marginal cells behind fore wing paranal cells: 2/3;
anx: fore wing 13/13, hind wing 9/8; pnx: fore wing
10/10, hind wing 10/10; cs under pt: fore wing 6/6,
hind wing 6/6; anal triangular cells: 4/5. Hind wing
31 mm. Abdomen 36 mm.
Venational details for paralectotype female: num-
ber of marginal cells behind fore wing paranal cells:
1/1; anx: fore wing 11/11, hind wing 9/9; pnx: fore
wing 9/9; hind wing 9/9; cs under pt: fore wing 5/4,
hind wing 4/5. Hind wing 29 mm. Abdomen 33
mm.
Variation. — Two teneral males of £. crotalinus
reared in Arizona are superficially similar to E. heter-
odon in the blunt shape of the cercus, the vestige of an
isolated antehumeral stripe in one specimen, and the
fuscous condition of the extensor surfaces of the met-
afemora. However, the penis structure of both speci-
mens is unmistakably that of £. crotalinus. Because
the two Arizona specimens are teneral, I cannot deter-
mine if dark thoracic stripes similar to those of £. het-
erodon would have developed, or whether the macula-
tion would have been the same as observed when
mature.
The male described by Hagen in Selys (1858) has
the darkest and most distinct antehumeral stripes that
I have seen for any specimen of this species. The tib-
iae of a majority of specimens I examined are entirely
dark brown with the lateral costae yellow; others, in-
cluding the lectotype, have the entire ventral margin
yellow.
Biology. — Specimens are apparently common at
favored localities. Boris Kondratieff (in litt, 1990)
states that adults of this species were ‘on every [ex-
posed] rock [in Rio Macoba] — hundreds and hun-
dreds of them.’ During a two-day stay at Ajijic near
Laguna Chapala, Oliver S. Flint, Jr., and M. A. Ortiz
collected 28 males and 23 females. Calvert (1907)
records the following concerning its habits:
‘[Progomphus borealis} and Erpetogomphus were found
on sand- or mud-banks, hardly above water-level,
along sides of an irrigating ditch into which the water
from the baths of Santa Rosalia empties. On alighting
on these banks, P. obscurus borealis held its abdomen
slanting upward to form an angle of 45°-60° with the
bank, while £. crotalinus held it nearly horizontal.’
The male collected at Lago de Patacuaro in
Michoacan is noted as taken on ‘stream flowing into
lake’, the two males and one female from the pyra-
mids at S. Juan Teotihuacan as ‘on high grassy plains
far from water. Gonzalez and Novelo (1991) record
GARRISON: Revision of Erpetogomphus
the following about £. crotalinus (in translation):
‘This is the most abundant gomphid in the area [the
Biosphere Reserve of La Michiliá, Durango]. Like
other members of the family, the adults of E. crotali-
nus have a seasonal flight pattern that, although it has
not been exactly established, apparently is restricted
to the months of May to October. The individuals of
this species are characteristic inhabitants of both sea-
sonal and permanent streams, easily recognized in the
field by the bright green colour of the thorax. Males
have the habit of resting on emergent rocks, a posi-
tion they defend aggressively from other conspecific
males.’
Erpetogomphus crotalinus has been collected with £.
designatus, E. lampropeltis and E. heterodon.
Collection dates range from May through October
(Calvert 1905).
Distribution (fig. 205). — Erpetogomphus crotalinus
is restricted to the higher xeric areas of central and
western Mexico. The species was reared from larvae
collected by Minter Westfall, Jr., from the Slaughter
Ranch, 15 mi W of Douglas, Cochise Co., Arizona;
but no other collectors have encountered it there. I
have also seen a pair from New Mexico (Eddy Co.,
Sitting Bull Falls), which may be strays from Mexico;
its occurrence in the southwestern United States ap-
pears sporadic, and other odonatists have failed to
find the species during their work in Arizona and
New Mexico. Altitudinal data show a range of 1225
m (Mexico: Chihuahua: Santa Rosalia Springs) to
2290 m (Mexico: Mexico: Pyramids, S. Juan
Teotihuacan). In the Paris Museum is a male labelled,
‘Veracruz’, which is probably in error.
Material
Type data. — Lectotype male by present designa-
tion with following data: white handwritten label by
Hagen, ‘G. lineatus | Mexico’; white handwritten la-
bel, ‘crotalinus in an unknown hand; white, black-
bordered printed label: ‘Mus. Berol.’; two small prin-
ted) labels Elagent;s red) labels wathipeiype
[printed]/12334’ (written by N. Banks); rectangular
label [all printed unless otherwise stated):
‘ERPETOGOMPHUS / CROTALINUS HAGEN [stamped]/
BB. @alvers, dee. 1905) [wetten NB ICHANEU END!
165 [written]. In mczc. The penis and hamules are
missing and probably served as the basis for the free-
hand illustrations of these structures for Hagen in
Selys (1858). Paralectotype female with following da-
ta: green printed label: ‘Mexico/ Deppe.’; red printed
label “Typus’; yellow printed label, “Zool. Mus./
Berlin’, on reverse side written in ink, ‘cat nr./ 2314;
white label: ‘syntypus 2 von: / Ophiogomphus | crot-
alinus Hagen’ [written in an unknown hand]; yellow
label (handwritten except as noted), PARALECTOTYPE
[printed] ®/ Ophiogomphus | crotalinus Hagen/ des.
229
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
[printed] R. W. Garrison 1984’. In zMHB.
There is a male in the ZMHB with the same data as
the paralectotype female, except as follows: green la-
bel, ‘Mexico Deppe’; green label, ‘crotalinus | Hag.’
[both handwritten by Selys]; printed label, ‘2314’;
and ‘Syntypus d von / Ophiogomphus | crotalinus
Hagen’. Although the specimen is labelled as a type, I
believe it is not a syntype and, in fact, that it repre-
sents the second male described by Hagen in Selys,
1858. I decided that the male in the mczc and the fe-
male in the ZMHB represent the original syntypes for
the following reasons: The original (1854) descrip-
tion of Ophiogomphus crotalinus is brief and was based
on a male and female. Hagen gave measurements for
both sexes (abdomen: male 36 mm, female 33 mm;
hind wing male 31 mm, female 29 mm). These meas-
urements and the description of the male (‘Front of
thorax yellow, with indication of two median stripes
and a pale red humeral stripe...’) correspond to the
MCZC male and the ZMHB female. Hagen im Selys
(1858) described this species in greater detail and had
access to a second, smaller male (abdomen 33 mm,
hind wing 29 mm). He stated, ‘In the other specimen
[z.e., the second male], the stripes are brown and
straight, quite visible...’ The original male is again
characterized as follows, “Thorax yellow green with-
out spots and...one can scarcely see the appearance of
two median stripes and a thick antehumeral stripe...’,
which tallies with the earlier description of 1854. The
male from the ZMHB corresponds with the smaller of
the two measurements given by Hagen in 1858; and
it also has a pair of easily visible antehumeral stripes,
an apparently rare feature for this species. The ZMHB
male is unique in this respect. Even though the origi-
nal description by Hagen gives ‘Musée de Berlin’ as
the repository of the types, I strongly suspect that
Hagen took the male with him when he was invited
to join the faculty at the Museum of Comparative
Zoology in 1867.
Other material (97d, 449, including lectotype d and
paralectotype 9). — U. S. A.: ARIZONA: Cochise Co., San
Bernardino Ranch, 15 mi E of Douglas (rsca); New
Mexico: Eddy Co., Sitting Bull Falls, Guadalupe Mtns.,
(RwG, wM); Mexico: Chihuahua: Chihuahua Creek, 4 mi
SW of Col. Juarez, (csuc); Rio Pacheco, (csuc); Santa
Rosalia Springs, 1219 m, (ANsP); Durango: Durango City,
(ansp); Rio Dalita and marshes, 6 mi E of Durango, 1890
m, (pre); Jalisco: Jopopau, 1667 m, (anse); Guadalajara,
(ANSP, MCZC); Laguna de Chapala (nr. Mex. Hwy 15), 75 mi
SE of Guadalajara, (RwG); Ajijic nr. Laguna Chapala (usnm,
cc, TWD); Ajijic, (rsca); Rio La Sanguijuela, Degollado,
(UNAM, RWG); route 15, km 206, San Lorenzo, 9 mi W
Tuxpan, (USNM, cc); Jalisco [no other locality], (BMNH);
Michoacan: Arrollo W of La Piedad, (UNAM); Lago de
Patzcuaro, 2.6 mi W of Quiroga, 2042 m, (rsca); Route 15,
km 206, San Lorenzo, 8 mi W of Tuxpan, (usnm); Route
15, km 431, Carapan, (UsNM); Route 15, km 291, near
Morelia, Parque Nacional Insurgente Morelos, (UsNM);
230
stream 0.5 mi S of Tuxpan, 1798 m, (pre); Hidalgo: ditch-
es around Tasquillo, 1700 m, (pre); Rio Tula at Puente
Tasquillo, (UsNM); Tecozautla, Rio San Juan, 1710 m,
(rwc); Tecozautla, Rio Tecozautla, 1750 m (rwG):
Tecozautla, arroyo San José del Desierto, 1730 m (rwG);
Mexico: Route 15, km 125, El Salitre, (usnm); Pyramids, S.
Juan Teotihuacan, 2290 m, (rsca, cuic); Morelos:
Cuernavaca (BMNH, ANSP, CUIC); Puebla: ditch just NW of
Tehuacan, 1700 m, (DRP, TWD); Veracruz: Veracruz [prob-
ably erroneous locality], (MNHP); Mexico [no other local-
ity], (ZMHB).
Erpetogomphus heterodon sp. n.
‘ (figs. 28-thx, 42, 43-abd, 65-hamules, 87-penis,
110, 111-app, 162-vertex, 170-leg, 193-vl, 206-
distr, 230-wings)
Type data. — Holotype male. U. S. A: New
Mexico: Catron Co., Tularosa River just E of Aragon
on N. Mex. Hwy 12, 28 July 1984 (R. W. and J. A.
Garrison). Allotype female. Same data as male. Both
in USNM.
Paratypes (364 , 22). — New Mexico: Catron Co.,
same data as holotype and allotype, 27-28 July 1984,
228, 19 (rwG); same data, 23 August 1964
(Clifford Johnson), 34, 12 (pre, rsca); Gila Cliff
Dwellings, 30 July 1983 (John E. Hafernik, Jr.), 4d
(RwG); Grant Co., Grapevine Campground, East
Fork of Gila River at N. Mex. Hwy 15, 5600 ft., 27
July 1984 (R. W. and J. A. Garrison), 116 (rwe);
Texas: Jeff Davis Co., Limpia Canyon in Davis Mtns.
State Park, 23 June 1958 (M. J. Westfall, Jr.), 14
(esca); Limpia Creek at Tex. Hwy 17, 3.4 mi NE of
Ft. Davis, 4500 ft., 24 August 1977 (Dennis Paulson,
Susan Hills), 16 (pre); Reeves Co., Balmorhea State
Park, 13 Sept. 1983 (S. W. Dunkle), 1d (swp);
Mexico: Chihuahua: 6 mi NNE of Boquilla, 16 July
1960 (Scheibner), 16 (RwG); Chihuahua Creek, 4 mi
S of Col. Judrez, 25 Aug. 1986 (B. C. Kondratieff),
1d (csuc); ‘Bosuchel [probably Basuchi]’, 18 Aug.
1950 (R. F. Smith), 28 (AMNH, RWG).
Description
Holotype male. — Entire face pale light green,
slightly darker on postclypeus and anterior of frons;
labrum, base of mandibles, labium ivory; vertex and
antennae black; base of antefrons with transverse fur-
row, tumid medially, laterally with a pair of pits; po-
stocellar tubercles small, no postocellar ridge; occiput
pale green, slightly tumid medially, hind margin frin-
ged with long hairs, crest with black along lateral
margin, slightly emarginate medially; postocciput gr-
een, transverse; rear of head dark brown, becoming
pale green laterally.
Prothorax light green with dark brown on middle
lobe extending laterally to pleura, two adjacent later-
odorsal small green spots on medial lobe. Synthorax
(fig. 23) light green with following dark brown: in-
verted wedge-shaped middorsal stripe, its ventral end
not touching collar, its dorsal end forming a narrow
stripe below antealar carina and meeting narrow hu-
meral stripe; small isolated antehumeral stripe; vesti-
gial second lateral stripe extending from metacoxa to
metaspiracle; upper end of third lateral stripe. Coxae,
trochanters ivory, femora ivory with black external
surfaces spreading to lateral areas at distal 0.25 (fig.
170), tibiae black with small lateral ivory stripes along
basal 0.5; tarsi and armature black.
Wings (fig. 230, paratype) hyaline, venation dark
brown, basal wing venation brown; anterior margin
of costa yellow; pterostigma brown, darker around
margin.
Venational details. Fifth antenodal thickened in all
wings; number of marginal cells behind fore wing
paranal cells: 2; anx: fore wing 14/12, hind wing 9/9;
pnx: fore wing 9/8, hind wing 10/10; cs under pteros-
tigma: fore wing 4/6, hind wing 6/4; anal triangular
cells: 4. Hind wing 33 mm.
Abdomen (fig. 42). Segment 1 pale orange-yellow
dorsally with narrow black lateral stripe, white lateral-
ly; segment 2 similar to segment 1 but with black
along anterior margin of segment laterally; segment 3
similar to segment 2 but with black along transverse
carina and posterior annulus; segments 4-7 similar to
segment 3 but black dorsolateral stripe incomplete
anteriorly so that dorsal orange-yellow merges with
lateral white and with black along transverse carina,
expanding ventrally, and with a similar mark posteri-
orly just before black annulus; segments 8-9 predom-
inantly yellow-orange with black denticles dorsally,
incomplete black dorsolateral stripe extending from
anterior margin of segment but ending just before
posterior margin; segment 10 yellow-orange posteri-
orly, orange-brown anteriorly. Abdomen 40 mm.
Cercus (fig. 110) yellow brown, strongly angulate
near middle, ventral margin with inferior carina at ba-
sal 0.30. Epiprocts curved at distal 0.75, tips dark-
ened, bluntly pointed, slightly divergent.
Accessory genitalia (fig. 65, paratype). Anterior ha-
mule black, divided at upper 0.30, lower branch form-
ing a small, truncate appendage; posterior hamule
white, triangular, with anterior basal shoulder, tip
with small, anteriorly recurved tooth; penis with small
serrated lateral lobes, cornuae (fig. 87, paratype) mod-
erately long, widely separated tips broadly rounded.
Allotype female. — Overall colouration similar to
male but with more extensive pale areas on abdomen
and legs; furrow anterior to medial ocellus as in male;
vertex (fig. 162) with postocellar tubercles not con-
nected but with small remnant of ridge extending
medially; occiput simple, posterior margin raised
with row of hairs; postocciput green, transverse.
Venational details. Fifth antenodal thickened in all
GARRISON: Revision of Erpetogomphus
wings; number of marginal cells behind fore wing
paranal cells: 3/2; anx: fore wing 13/12, hind wing
10/10; pnx: fore wing 7/8, hind wing 9/10; cs under
pterostigma: fore wing 6/6, hind wing 6/6. Hind
wing 36 mm.
Abdomen (fig. 43) as in male, but with black dor-
solateral stripe reduced, with black extending anteri-
orly before transverse carina. Abdomen 37 mm.
Vulvar lamina (fig. 193) with broadly planar rec-
tangular flaps meeting at anterior margin; juncture of
Y-shaped postlamellar ridge at level of posterior mar-
gin of lamina.
Diagnosis
This species is related to E. crotalinus and E. sipe-
don. Erpetogomphus crotalinus lacks the dark thoracic
stripes present in E. heterodon (fig. 22); the extensor
surfaces of the tibiae of Æ. heterodon are black (fig.
170) (yellow in E. crotalinus); and the cornuae of the
penis in E. heterodon are broadly rounded (fig. 87)
(long and acutely pointed in £. crotalinus [fig. 86]).
Males of E. heterodon are most similar to E. sipedon,
but they differ in thoracic maculation shape of the
epiprocts, and structure of the penis. In E. sipedon,
the antehumeral stripe is long and joined with the hu-
meral stripe (fig. 19); in £. heterodon, this stripe is ves-
tigial and disjunct. In £. sipedon, the tips of the super-
ior appendages are slender and acuminate (figs.
105-106), not obtusely acuminate as in E. heterodon
(figs. 110-111). The cornua of the penis in £. sipedon
(fig. 83) possess a mesal lobe absent in Æ. heterodon
(fig. 87).
Structurally, the females of E. crotalinus, E. hetero-
don, and E. sipedon approach one another closely.
The latter two species have a relatively narrow occiput
with the width across the crest about 4 times as long
as its greatest length. The occiput in £. crotalinus is
wider, about 3 times the width. I have been unable to
detect morphological differences between females of
E. heterodon and E. sipedon. Body maculation must be
used to separate females.
Remarks
Variation. — Slight variation occurs in males within
the type series. The dark dorsolateral stripe on abdo-
minal segment 3 may be broken in the middle. In the
male from Mexico (6 mi NNE Boquilla), this stripe is
reduced to an isolated spot on the basal 0.30 of the
segment. Similarly, the extent of black markings on
abdominal segments 8-10 may be reduced.
Venational statistics for type series. Males: Number
of marginal cells behind fore wing paranal cells: 0-3;
anx: fore wing 11-14/11-15, hind wing 8-11/8-11;
pnx: fore wing 7-11/7-11, hind wing 8-12/8-11; cs
under pterostigma: fore wing 4-7/4-6, hind wing 4-
7/4-7; anal triangular cells: 3-4. Hind wing 32-34
231
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
mm. Abdomen 36-40 mm.
Females: Number of marginal cells behind fore
wing paranal cells: 1-4; anx: fore wing 13/12-13, hind
wing 8-10/9-10; pnx: fore wing 7-9/8-9, hind wing
8-10/8-10; cs under pterostigma: fore wing 5-6/5-6,
hind wing 5-6/5-6. Hind wing 34-36 mm. Abdomen
36-37 mm.
Biology. — Adult males were taken along sandy
margins of the Tularosa River in New Mexico. They
often sat on the bank facing the river and were easy to
approach. I collected one female as it hovered over
the river ovipositing. Another teneral female was
flushed from nearby vegetation. At the East Fork of -
the Gila River, males sat on emergent rocks in the
stream or by the river’s edge. They occasionally made
swift sorties, usually to chase another male. At this lo-
cale, E. heterodon was sympatric with £. lampropeltis
natrix. Altitudinal data show E. heterodon to occupy
high, mostly xeric areas of the southwestern United
States and northern Mexico. Collection dates are
from 23 June to 13 September at elevations of 1370
m (Limpia Creek, Texas) to 1700 m (Grapevine
Campground, New Mexico).
Distribution (fig. 206). — Erpetogomphus heterodon
is known only from western Texas, New Mexico, and
northern Mexico. It has been collected with E. crotal-
inus in at least one locality in Mexico (Chihuahua
state, Rio Pacheco, 27 Aug. 1986, B. Kondratieff, S.
Dunkle, pers. comm.), and is apparently allopatric
with E. sipedon.
Erpetogomphus compositus Hagen in Selys
(figs. 24, 25-thx, 66, 67-hamules, 88, 89-penis, 112,
113-app, 145-ept, 139-base of postfrons, 163-vertex,
194-vl, 206-distr)
Erpetogomphus compositus Hagen in Selys, 1858: 660 (400
sep.) (9, ‘Rivière Peros [sic Pecos] ( Texas occidental)’). —
Hagen in Selys 1859: 536 (10 sep.) (descr. 2); Hagen in
Selys 1873a: 740 (12 sep.) (descr. d, 9, ‘Le nord de la
Californie”); Selys 1873b: 519 (75 sep.) (list); Calvert
1905: 166 (Ariz.); Calvert 1908a: 45 (Ariz.); Muttkowski
1910: 86 (cat.); Calvert 1912a: 289 (mentioned);
Williamson 1914a: 226 (Ariz.); Kennedy 1917a: 544
(Calif.); Seemann 1927: 22 (Calif.); Byers 1928: 51 (lar-
va unknown); Needham and Heywood 1929: 80 (descr.);
Williamson and Williamson 1930: 12 (summary of stat-
us); Tinkham 1934: 215 (W. Tex.); Ahrens 1938: 11
(Ariz., Utah); Fraser 1940: pl. 5 (penis); Ferguson 1940:
5 (Tex.); Needham and Westfall 1955: 144 (descr.);
Pritchard and Smith 1956: 116 (key); Gloyd 1958: 8 (W.
Tex.); Musser 1961: 54 (larva, Utah); Musser 1962: 14
(larva, Utah); Borror 1963: 104 (common name);
Cruden 1964: 81 (Calif.); Montgomery 1968: 133
(distr.); Paulson and Garrison 1977: 151 (Washington);
Molnar and Lavigne 1979: 130 (Wyo.); Paulson 1982:
255 (distr.); Paulson 1983: 67 (Wash.); Davies and
Tobin 1985: 27 (cat.); Tsuda 1986: 87 (cat., attributes
authorship to Selys); Bick 1990: 3 (Idaho); Bridges 1991:
232
VII: 49 (cat.); Tsuda 1991: 5 (cat.).
Gomphus compositus. — Hagen 1861: 99 (desc. 9, Tex.).
Gomphus (Herpetogomphus)? viperinus. — Hagen, 1873 (mis-
identification, Yellowstone).
Herpetogomphus compositu. — Hagen 1874: 597
(Yellowstone, Tex.); Hagen 1875a: 42 (Tex.); Hagen
1875b: 918 (N. Mex.); Selys 1879: 64 (2 sep.) (characters
of genus); Kirby 1890: 60 (cat., attributes authorship to
Selys); Calvert 1899: 386 (characters of genus); Currie
1903: 303 (Ariz.); Osburn 1905: 186 (mentioned); Byers
1928: 5 (larva unknown); La Rivers 1938: 85 (Nev.); La
Rivers 1940a: 112 (Nev.); La Rivers 1941: 177 (Nev.).
Erpetogomphus coluber Williamson and Williamson, 1930:
17 (d, San José de Comandu, Baja Calif. Sur). —
Needham and Westfall 1955: 143 (desc.); Borror 1963:
104 (common name); Montgomery 1968: 133 (distr.);
Montgomery 1973: 239 (comment on name); Paulson
1982: 255 (distr.), 266 (synonymy with E. compositus);
Bridges 1991: VII: 49 (cat.).
Description
Male. — Entire face pale grey green with dark mar-
kings as follows: along basal margin of labium and
median extension ending before distal margin (almost
absent in some specimens), frontoclypeal suture, base
of antefrons; vertex all dark brown or with pale me-
dian area extending posteriorly from median ocellus
to anterior margin of occiput; antennae dark brown;
anterior margin of vertex forming a trough at junc-
tion of postfrons, this trough with a pair of deeper
pits anterolateral to median ocellus (fig. 139); occiput
white green, wide, planar with slight tumid median
area, crest barely convex, slightly to broadly emargi-
nate medially, its hind margin covered with pale
brown hairs; postocciput white green, transverse, not
visible dorsally, rear of head dark brown.
Prothorax pale green with brown dorsolaterally on
middle lobe; synthorax (fig. 24) predominantly pale
green with following dark brown: well defined mid-
dorsal stripe widening to collar, its upper end extend-
ing along antealar sinus connecting with well defined
humeral stripe; antehumeral stripe widened dorsally
and connected basally at mesinfraepisternum; lower
part of humeral stripe extending posteroventrally but
not connecting with well defined sinuate second lat-
eral stripe; lower 0.50 of this stripe encompassing
metaspiracle, constricted along posterior margin at
0.50 and abruptly widening dorsally before turning
anterodorsally toward subalar carina; complete third
lateral stripe ending behind posterior margin of met-
acoxa. Pale colour of thorax light green with lighter
tone almost becoming white in areas between antehu-
meral and humeral stripes, and between second and
third lateral stripes. Coxae, trochanters white grey
with slight wash of brown, tibiae pale grey green with
defined superior surfaces black, basal 0.40 of metafe-
mora with streaks of grey green or with basal 0.60 of
this area largely pale, tibiae black with narrow stripe
of pale grey green along basal 0.30 of lateral margin in
some specimens; tarsi and armature black.
Wings hyaline with slight wash of yellow at ex-
treme base (more so on teneral specimens), basal wing
venation brown, especially anteriorly, anterior margin
of costa yellow white except for anterior margin of
pterostigma, costa beyond pterostigma white or dark-
ened, but contrasting with black of pterostigma; pte-
rostigma black.
Venational statistics. Fifth antenodal thickened in
all wings; marginal cells behind fore wing paranal
cells: 0-4; ; anx: fore wing 10-15/10-15, hind wing 7-
11/7-11; pnx: fore wing 6-11/6-10, hind wing 7-
11/7-11; cs under pterostigma: fore wing 3-6/3-6,
hind wing 3-6/3-6; anal triangular cells: 2-5. Hind
wing 25-32 mm.
Abdomen. Segment 1 pale grey green with broad
basal semicircle of brown laterally, its posterior end
touching lateral articulated area; segment 2 white
with dorsolateral brown stripe sending a ventral stem
posterior to auricle, and connecting with black annu-
lus, a small brown spot at anteroventral margin of
segment; segment 3 with isolated black dorsolateral
spot surrounding lateral carina, distal 0.25 of segment
with dorsolateral black increasing in width posterior-
ly and touching dorsally near black annulus; segment
4 similar to segment 3, but isolated black of transverse
carina longer and touching or connecting with distal
black, thus forming an incomplete midlateral stripe
occupying distal 0.75 of segment and isolating ventral
longitudinal wedge of white; segments 5 and 6 like
segment 4, but with middorsal white becoming fus-
cous; segment 7 with anterior 0.5 white, except for
tawny transverse carina, becoming entirely tawny
posteriorly, some specimens with varying degrees of
lateral black on posterior 0.25 of segment or in speci-
mens from Baja California (£. coluber) with tawny ar-
eas becoming black; segments 8-10 yellow brown
with darker red brown dorsally, in Baja California
specimens this dark red brown replaced with well de-
fined black on segments 8 and 9, and dark brown on
segment 10. Abdomen 32-39 mm.
Cercus (figs. 112-113) ocher, in lateral view slight-
ly concave dorsally near base and with tips slightly re-
cumbent, cercus enlarged ventrally at basal 0.45 with
inferior carina along this same area, remainder slight-
ly concave with tip forming a blunt point. Epiprocts
yellow brown, gently curved dorsally at 90°, tips (fig.
134) dorsoventrally flattened and truncate or
obliquely truncate.
Accessory genitalia (figs. 66-67). Anterior hamule
dark brown, divided at basal 0.5; lower branch form-
ing a small, posteriorly directed truncate appendage;
posterior hamule grey white, triangular; with anterior
basal shoulder, tip bluntly rounded and armed with a
small anteriorly directed tooth on posterior border;
penis with lateral lobe small, roughly semicircular,
GARRISON: Revision of Erpetogomphus
serrate; cornuae (figs. 88-89) moderately long, widely
separated, tips broadly rounded.
Female. — Head as in male but pale areas more ex-
tensive, especially on vertex, where dark brown may
be reduced to areas around ocelli and postocellar pro-
tuberances; occiput narrower than in male, with erect
curvilinear occipital crest separating easily visible
postocciput (fig. 163), posterior margin of postocci-
put slightly concave; prothorax and thorax as in male,
but with pale areas more extensive; wings with yellow
at base more extensive, often with markings within
second series of antenodals to level of arculus and cu-
bitoanal area in both wings.
Abdomen in well preserved specimens with pale
colours primarily white with pale orange dorsally,
black dorsolateral stripes reduced and separated in
their middle on segments 3-6; segment 7 similar to
male but with posterior 0.25-0.30 darker laterodor-
sally; segments 8-10 tawny, with varying amounts of
dark brown, especially dorsally, on segments 8-9; cer-
ci ocher. Abdomen 31-37 mm.
Vulvar lamina (fig. 194) with prominent basal
plates, quadrangular laminar plates acuminate poste-
riorly, meeting at anteromedial margin; cleft between
plates forming a 90° angle; stem of Y-shaped postla-
mellar ridge at level of posterior margin of lamina.
Venational statistics: number of marginal cells be-
hind fore wing paranal cells: 1-6/0-5; anx: fore wing
12-15/12-14, hind wing 8-11/9-11; pnx: fore wing 7-
11/8-10, hind wing 9-11/8-11; cs under pterostigma:
fore wing 3-5/4-6, hind wing. Hind wing 30-32 mm.
Diagnosis
This common, distinctive species is easily separated
from its nearest ally, E. lampropeltis, by several struc-
tural and maculation characters. Males of E. composi-
tus differ from E. lampropeltis in lacking the characte-
ristic middorsal angulation of the cercus. The tip of
the epiproct in E. compositus is truncate (fig. 134), but
this structure in Æ. lampropeltis is obliquely and
bluntly pointed (fig. 132). Depressions in male and
female anterolateral to median ocellus will also sepa-
rate the species: in E. compositus, an arcuate trough
has more pronounced pits anterolateral to the median
ocellus (fig. 139). In £. lampropeltis, the anterolateral
pits are deeper and the intervening area directly ante-
rior to the median ocellus is not as depressed (fig.
138). This structure in £. lampropeltis more closely
approaches the condition found in £. viperinus (fig.
37):
Erpetogomphus compositus, though a boldly marked
species, is lighter than £. lampropeltis. The occipital
crest in E. compositus is largely pale, though there may
be some darkening along the lateral 0.30 of margins
in specimens from Baja California. In E. lampropeltis,
the occipital crest is entirely dark brown and provides
239
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
a striking contrast to the pale occiput. The thoracic
patterns of the two species are different (figs. 20-21,
24-25), and the dorsolateral black stripes of abdomi-
nal segments 4-6 of E. lampropeltis are more exten-
sive.
In the field, pale thoracic colouration will easily
separate both species. Erpetogomphus compositus is
unique in having a pale green thorax with intervening
grey white areas between the antehumeral and hu-
meral stripes and second and third lateral stripes. In
E. lampropeltis, the thoracic colouration is grey green
(E. lampropeltis lampropeltis) or dark green (E. lam-
propeltis natrix).
Females of E. compositus differ from E. lampropeltis
in the shape of the occiput. In Æ. compositus, the occi-
put is narrow and the postocciput is clearly visible in
dorsal view (fig. 163). In E. lampropeltis, the occiput
is broad and the postocciput is not visible dorsally
(fig. 160). Secondly, the area in front of the median
ocellus mirrors the condition stated for males above.
The vulvar laminae of the two species appear indistin-
guishable.
Remarks
Williamson and Williamson (1930) described in
detail Erpetogomphus coluber, comprising 31 males fr-
om San José de Comandu, Baja California. In com-
paring E. coluber with other species, they stated that
E. coluber would, in Calvert’s (1905) key to the genus
“run to AA, page 160, and if the individual be one
with the facial dark markings reduced, it will run to
H. compositus. Drs. Calvert and Kennedy regard colu-
ber as distinct from any described species.’ The
Williamsons sent two males to Calvert, who com-
mented, ‘No special reason for thinking it diadophis
[= E. eutainia); like a small compositus, differs in size;
has broader metepisternal dark stripe; brown on [ab-
dominal segments] 3-7 more extended; brown on 8-
10 much darker; darker lines or stripes on the fronto-
clypeal and clypeo-labral sutures’. Finally, the
Williamsons commented on the affinities of E. colu-
ber to E. compositus as follows: “Compositus is certain-
ly its closest relative and the derivation of coluber fr-
om compositus, through geographical isolation in Baja
California, is almost certain and is a case exactly pa-
rallel, so far as speciation goes, to that of certain spe-
cies, of several genera, endemic in Florida. Coluber is
separated from compositus by its darker color, espe-
cially of the last four abdominal segments and by ve-
national characters, especially the two-celled anal
triangle and the single row of cells posterior to A in
the front wing.’
I have seen the holotype and 15 paratypes of E. co-
luber and believe, as did Paulson (1982), that they re-
present diminutive, melanic examples of E. composi-
tus. The specific differences ascribed to E. coluber are
234
those of venation only, and I believe these to be cor-
related to their smaller size. I have found no differen-
ces in body morphology, including the accessory ge-
nitalia.
Variation. — Two specimens from San Diego
County appear intermediate to E. coluberand E. com-
positus s.s. These males have more pronounced dark
thoracic stripes, but the dorsum of abdominal seg-
ments 8-10 lacks dark brown markings typical of £.
coluber. However, other E. compositus have varying
amounts of dark brown on these segments. The hind
wing anal triangle in 3 of 16 specimens of £. coluber
. has 3 anal triangular cells instead of 2, and a 2-celled
anal triangle, though rare, does exist in £. compositus
(s.s.). For example, of 21 males I measured from
California, Arizona, and another from northern Baja
California, one of the San Diego males and another
from Riverside, California, had one wing each with
two cells. These two specimens had shorter hind
wings (26 mm and 28 mm, respectively) than is typi-
cal for E. compositus (s.s.). Statistics for the number of
anal triangular cells show it to be variable within £.
compositus (s.s.). The number of wings with range of
stated conditions is (number of wings follows in pa-
rentheses): 3 (19), 4 (19), 5 (2). Hind wing lengths
from the San Diego males are 25-26 mm, well within
the range for E. coluber. The next smallest specimen is
a male from Riverside (28 mm), the same male with
two anal triangular cells in one hind wing.
Most wings of E. coluber which I have examined
have no marginal cells behind the fore wing paranal
cells, but 6 of 32 (19%) did. Eight of 42 (19%) wings
of E. compositus (s.s.) had no marginal cells.
Williamson and Williamson (1930) warned of fal-
libility of some of their characters among the para-
types they examined: “The face markings are very pro-
nounced in some and scarcely discernible in others;
there is considerable variation in the extent of the
dark thoracic markings [fig. 25] and this variation oc-
curs independently on the mesepisternum and mete-
pisternum, so an individual may have the dark hu-
meral and antehumeral relatively extensive and the
two dark lateral stripes relatively reduced and vice ver-
sa.
With the exception of specimens from Baja
California and the two specimens from San Diego,
body colouration appears remarkably constant
throughout its range. The male from Baja California
Norte (Guadalupe Hot Springs) is more like typical
E. compositus, but it has no marginal cells behind the
fore wing paranal cells. The hind wing length (29
mm) and other venational characters indicate a closer
relationship to more northerly populations.
The female holotype was collected with the type se-
ries of E. designatus and was probably collected in the
vicinity of Roswell, New Mexico in 1854-1855. See
remarks under £. designatus for further details.
Venational details. Holotype female of E. composit-
us. number of marginal cells behind fore wing paranal
cells: 2/4; anx: fore wing 13/13, hind wing 9/10; pnx:
fore wing 8/9, hind wing 9/8; cs under pterostigma:
fore wing 5/6, hind wing 5/6. Holotype male of £. co-
Luber. number of marginal cells behind fore wing par-
anal cells: 0/0; anx: fore wing 12/12, hind wing 9/9;
pnx: fore wing 7/7, hind wing 9/9; cs under pterostig-
ma: fore wing 2/2, hind wing 4/4.
Biology. — This species and Progomphus borealis
McLachlan in Selys are the two most conspicuous
gomphid elements along most desert streams and irri-
gation ditches in the southwestern United States. In
Arizona, this species occurs commonly in the Lower
Sonoran Zone, and in ecotonal fingers into the Upper
Sonoran Zone. Kennedy (1917a), who collected £.
compositus at certain sites in the northern Central
Valley of California, writes: ‘One female, a teneral,
was taken on the irrigating ditch across the river from
Oroville.’ Collecting along the lower Truckee River
south of Pyramid Lake, Nevada, Kennedy (1917a)
mentions that ‘This widely spread species occurred
sparingly on the riffles of the lower Truckee. The
males appeared to be more nervous and more touchy,
flying farther for conflict with passing males than the
males of Ophiogomphus morrisoni nevadensis.
The two males collected at the Boyce Thompson
Southwestern Arboretum in Pinal County, Arizona,
had the following annotation by H. K. Gloyd: ‘[in]
Creek, arboretum. Canyon, after rain and run-off al-
most over. between 5 & 7: 00 p.m., water low, few
dragonflies seen’. Williamson and Williamson (1930)
provided a lengthy account of the type locality and as-
semblage of Odonata present with Erpetogomphus co-
luber. In August, 1972, I collected several £. compos-
itus with Stylurus plagiatus (Selys) at Riverside Park,
Yuma, Arizona. Adults were taken in the late after-
noon on grassy areas shaded by trees. Adults of this
species and Progomphus borealis were abundant along
Big Chico Creek in Chico, Butte County, California
in June and July of 1974, 1976, and 1978. They sat
on exposed sand bars near the edge of the creek, and
both were difficult to approach. Progomphus borealis
was the more wary. In the late afternoon, I have col-
lected £. compositus away from water, resting on dry
desert scrub. Collection dates range from 24 May
(Nevada: Clark Co.) to 13 Dec. (Calif: San
Bernardino Co.).
Distribution (fig. 206). — Erpetogomphus composit-
us is a desert species found commonly in the south-
western United States. It has been taken as far east as
Dallas (Hagen, 1875a), but most records from Texas
are from the western portion of the state. It occurs in
eastern New Mexico and is common in low desert re-
gions of Arizona, especially southwest of the
GARRISON: Revision of Erpetogomphus
Mogollon Rim. It is common in the arid regions of
southern California and penetrates north through the
Central Valley to Chico. Paulson and Garrison
(1977) list it from south central Washington. The
species was listed from Oregon by Hagen (1875a,
1875b) with no specific locality); but specimens were
recently collected from the John Day River,
Deschutes Co. Oregon (Valley 1993).
Erpetogomphus compositus occupies river systems in
the Great Basin of Nevada, but its most easterly dis-
tribution is unknown. Bick (1990) lists a specimen
from Owyhee Co., Idaho. Hagen (1874) records it
from the Yellowstone, but it was probably taken in a
more arid environment around what is present-day
Yellowstone National Park. Molnar and Lavigne
(1979) also cite it without specific locality from
Wyoming, based on previous records given by
Needham and Heywood (1929), Needham and
Westfall (1955), and Pritchard and Smith (1956). All
of these records probably originated from the original
Hagen (1874) citation. In Utah, the species apparent-
ly penetrates only the southwesternmost part of the
state. The most southerly distribution for £. compos-
itus is also unknown. Except for E. coluber, the only
records I have observed from mainland Mexico are
two males collected by J. H. Williamson in
Hermosillo, Sonora. The distribution of E. composit-
us in Texas, New Mexico, and Arizona indicates that
it must be further widespread in northern Mexico.
Material
Type data. — Of E. compositus. Holotype female
with following label data: ‘Aug/ 16° [handwritten],
‘Hagen’ printed [two of these labels attached], ‘com-
positus written in an unknown hand; red label ‘Ho-
LOTYPE [printed] / Erpetogomphus compositus | Hagen
1857 [sic, should be 1858] ‘9’ handwritten by rwc.
In mczc. Of E. coluber. Holotype male dry in envelo-
pe with following data: ‘E. coluber written in pencil
by E. B. Williamson, ‘Mexico/ State of Baja
California/ San Jose de Comandu/ J. H. Williamson/
Oct 10 1923 [all stamped]/ 8214.” handwritten by
EBW, ‘Type handwritten in pencil by EBW on side
of label. In ummz.
Type locality of £. compositus herein restricted to
vicinity of Roswell, Chaves Co., New Mexico (see re-
marks under £. designatus).
Other material (1516, 469, including holotype 2 E.
compositus and holotype 3 E. coluber). — U. S. A.: ARIZONA:
Cochise Co., Miller Cyn., Huachuca Mtns., (cc); pond at
Slaughter Ranch, San Bernardino Valley, 15 mi E of
Douglas, 1169 m, (rw6); 5 mi E of Hereford (cc); San
Pedro River, 9 mi SW of Tombstone, (Byuc); San Pedro
River, at Hereford Rd., (rw); Coconino Co., Havasu
Canyon, 3 mi N of Supai, 923m, (RWG); Supai, (CDFA);
Graham Co., 3 mi SE of Bylas, valley of the Gila, 2000 ft.,
(ummz); Roper Lake, 5 mi S of Safford (rwe); Maricopa
235)
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
Co., Granite Reef Dam, (ummz); Mesa, (ummz); Phoenix,
(rw); slough ponds by Verde River, by Ariz. Hwy 87, Ft.
McDowell Indian Reservation (RWG); Tempe (UMMZ);
Mohave Co., spring, Hwy 15 bridge, Littlefield, (Byuc,
Rwe); Virgin River, Big Bend, (Byuc); Pima Co., Organ
Pipe Cactus Nat'l. Mon., Quitobaquito, (Long Beach State
Univ.); Quitobaquito Springs, 15 mi S of Lukeville, Organ
Pipe Cactus Natl. Mon., (rwG); Pinal Co, Boyce
Thompson Southwestern Arboretum, 4 1/2 mi SW of
Superior, (ummz); 8 mi NW of Florence, (ummz); Santa
Cruz Co., Santa Cruz River, Sonoita Creek Ranch (on way
to Salero Cyn.), ca. 2 mi W of Patagonia, by Ariz. Hwy 82,
(RWG); Yavapai Co, Oak Creek at Cornville, 1077 m,
(csuc); Yuma Co. Ave 3E at Co. 14 St., SE of Yuma, (RWG);
Riverside Park, nr. Colorado River, Yuma, (RwG); N. R.
Adair Park, McPhaul Bridge, by Gila River, ca. 15 mi ENE
of Yuma, (RWG); CALIFORNIA: Butte Co., Bidwell City Park,
by Big Chico Creek, Chico, 61 m, (Rwe); Oroville, (ummz);
Fresno Co., Friant, San Joaquin River, (ummz); Imperial Co.
Calexico, (Byuc); canal 8 mi E of Holtville, sea level, (DRP);
ditch 4.7 mi E of Bond’s Corner, (DRP); Hot Mineral,
(LACM); irrigation canal 7.3 mi E of Holtville, (pre); irriga-
tion canal at Winterhaven, (DRP); Inyo Co., Hunter Cyn.,
Salino Valley, (Lacm); Laws, Owens River, (ummz); Lone
Pine, Owens River, (ummz); Los Angeles Co., Tanbark Flat,
(cpra); Mono Co., 5 mi N of Benton Station, 5300 ft,
(UMMZ, LACM); Riverside Co. Blythe, (cc); Coachella Valley
Preserve, NE of Thousand Palms, (J. Cole); Palm Springs,
(LAcM); Whitewater River nr. Salton Sea, (DRP); San
Bernardino Co., Colorado River at Moabi Rd., S of Needles,
(RwG); Lost Palm Cyn., Joshua Tree N. Mon., (Long Beach
State Univ.); Parker Dam, (rwe); San Diego Co: T14S,
RSE, (casc); San Felipe Creek, 13.8 mi E of Julian, 615 m,
(pre); San Felipe Creek at Scissors Crossing, 11.9 mi E of
Julian, 738 m, (DRP, RwG); Sentenac Cyn., (LACM);
Vallecito, (LACM); Yolo Co., Cache Creek at William H.
‘Bill’ Davis Memorial Picnic Area, by Calif. Hwy 16, 6 mi N
of Rumsey, 132 m, (RwWG); Nevapa: Clark Co., Logandale,
(Byuc); Elko Co., Carlin, Humboldt River, (ummz); 8.5 mi
NW of Currie, (ummz); Humboldt Co., Can Spring, 3 mi S
of Pahuhe Meadows, (ummz); Lincoln Co., 6 mi S of Alamo,
(ummz); Nye Co., Amargosa R., 2 mi below Beatty, (ummz);
Pershing Co., Lovelock, Humboldt River, (ummz); Washoe
Co., Pyramid Lake, Truckee River, (ummz); White Pine Co.,
2 mi NW of Preston, (ummz); Texas: Brewster Co., Big
Bend Natl Park, 1 mi N of Rio Grande Village, (rwe);
Boquillas Cyn., Big Bend Natl. Park, (csuc); Rio Grande
Village, Big Bend Natl. Park, (csuc); Maverick Co.,
Quemado, (Lacm); UTAH: Washington Co., St. George, (cc);
Beaver Dam Wash, Terry Ranch, (Byuc); Beaver Dam
Wash, Lytle Ranch, (Byuc); Gunlock, W of Veyo, (Byuc);
Virgin River, Virgin, (Byuc); Mexico: Baja Calif: Norte.
Guadalupe Hot Spgs., Guadalupe Cyn., Sierra Juarez,
(csuc); Baja Calif: Sur. San José de Comandu, 10 Oct. 1923
J. H. Williamson), 156 (all paratypes of E. coluber) (ummz,
RWG, FSCA, USNM); Sonora: Hermosillo, (UMMZ).
Erpetogomphus boa Selys
(figs. 26-thx, 44, 45-abd, 68-hamules, 90-penis,
114, 115-app, 135-ept, 164-vertex, 167-
postocciput, 195, 196-vl, 207-distr)
Erpetogomphus boa Selys, 1859: 37 (11, sep.) (descr. d ‘Vera
Cruz, Mexique. Par M. Salle. (Collect. Selys.)’). — Walsh
1863: 253 (mentioned); Selys 1873b: 519 (75, sep.) (list);
236
Calvert 1905: 165 (notes); Calvert 1907: 399 (possible
identity with E. elaps); Muttkowski 1910: 86 (cat.); Ris
1917: 153 (notes and descr. of 2nd known gd);
Williamson and Williamson 1930: 11 (summary of stat-
us); Paulson 1982: 255 (Mex.); Davies and Tobin 1985:
27 (cat); Tsuda 1986: 87 (cat); Bridges 1991: VII.31
(cat.); Tsuda 1991: 95 (cat.).
Erpetogomphus crotalinus, nec Hagen in Selys, 1854. — Selys
1859: 537 (11, sep.) (2 descr. “Vera Cruz, Mexique. Par
M. Salle. (Collect. Selys.)’ = E. crotalinus Hagen in Selys,
1858.
Gomphus boa. — Hagen 1861: 100 (descr. from Selys).
Herpetogomphus boa. — Walsh 1862: 389 (mentioned);
Hagen 1875a: 42 (cat.); Selys 1879: 64 (2, sep.); Calvert
1899: 386 (list, English translation of Selys, 1879);
Calvert 1908b: 693 (mentioned).
Description
Male. — Labrum, clypeus, and frons light blue gr-
een, paler along sides of labrum and lateral lobes of
postclypeus; base of mandibles pale green, tips black;
base of antefrons anterior to median ocellus with nar-
row, arcuate, V-shaped sulcus, shallower medially;
vertex dark brown with usual postocellar tubercles,
occiput tumid medially, pale green brown, its posteri-
or margin ciliated, transverse to slightly concave,
slightly emarginate in the middle, or more rarely
smoothly curved, postocciput green, tumid; rear of
head red brown.
Prothorax red brown, synthorax (fig. 26) predomi-
nantly green, area around middorsal carina washed
with brown, often appearing as a faded middorsal
stripe; antehumeral and humeral stripes dark brown,
united into one large stripe; most of this stripe occu-
pying posterior 0.30 of mesepisternum, metastigma
black, a wash of brown between metepisternum and
metepimeron often forming an obscure, narrow tho-
racic stripe, subalar carina darkened with brown, me-
sinfraepisternum and metinfraepisternum brown;
metasternum green-brown. Coxae pale grey- brown
with some green on exterior surfaces, trochanters grey
brown, femora tawny at base, becoming dark brown
distally, tibiae and tarsi black.
Venational statistics (n = 13, including lectotype
3). Fifth antenodal (occasionally fourth, sixth, or
seventh) thickened in all wings; marginal cells behind
fore wing paranal cells: 0- 4/0-4; anx: fore wing 13-
15/13-16, hind wing 9-11/10-12; pnx: fore wing 9-
12/9-11, hind wing 10-13/10-13; cs under pterostig-
ma: fore wing 5-7/5-7, hind wing 5-7/5-7; anal
triangular cells: 4-5. Hind wing 33-34 mm.
Abdomen (fig. 44). Segment 1 brown dorsally, be-
coming green laterally; segment 2 with blue green
middorsal stripe, with dorsolateral stripe of brown
surrounding green auricle, lateral margin near ha-
mules pale greenish white; segment 3 with green mid-
dorsal stripe bounded on sides by dorsolateral stripes
of brown, lateral areas of tergites pale green becoming
white posteriorly, anterior transverse carina edged
with black expanding laterally interrupting white lat-
eral areas of segment, thus forming two pale spots,
posterior transverse carina also black, expanding later-
ally so that posterior 0.2 of segment is black; segments
4-6 similar to segment 3, but with middorsal brown
stripe narrowed anteriorly so that whitish green side
almost touches middorsal green; segment 7 with ante-
rior 0.5 dull white, conspicuously traversed by black
anterior transverse carina, posterior 0.5 of segment
red brown; segments 8-10 red brown with ventral and
posterior borders of each segment black. Appendages
red brown with apices becoming black. Abdomen 37-
40 mm.
Cercus (fig. 115) nearly straight, swollen anterior-
ly, apical 0.5 abruptly concave dorsally and covered
with thick series of strong black bristles. Epiprocts
about 0.5 the length of superiors, strongly curved
dorsally; tips slightly divergent, each a thick, blunt
point (fig. 135).
Accessory genitalia (fig. 68). Anterior hamule
black, glabrous, deeply forked with posterior branch
slightly longer than anterior, the whole structure re-
sembling a talon; anterior branch of hamule with a
small semi-oval planar surface on outer side, this area
covered with small hairs. Posterior hamule pale green
or white, becoming brown near tip; in lateral view, tip
obtusely pointed with posterior margin gently con-
vex. Penis (fig. 90) with lateral lobes poorly developed
posteriorly, almost circular, its margin serrated.
Female. — Overall colouration as in male, but with
pale areas on abdomen and legs more extensive. On
abdomen (fig. 45), brown on terminal segments ex-
tending anteriorly so that green middorsal and dorso-
lateral brown stripes become ill-defined. Lateral mar-
gins of abdominal segments mostly white and similar
to male except that white extends posteriorly to seg-
ment 8. abdomen 37-40 mm.
Vertex with slightly tumid, oval areas posterior to
lateral ocelli; posterior margin of occiput (fig. 164) el-
evated and nearly straight; postocciput green, tumid
medially (fig. 167).
Venational statistics. Fifth antenodal (occasionally
fourth, sixth, or seventh) thickened in all wings; anx:
fore wing 13-16/13-16, hind wing 10-12/10-11; pnx:
fore wing 9-13/9-12, hind wing 10-13/10-14; cs
under pt: fore wing 5-7/5-8, hind wing 5-8/5-8.
Hind wing 34-37 mm.
Vulvar lamina (fig. 195) membranous, diagonally
corrugated, outer margin of plate with strong costate
ridge, area immediately mesal to ridge furrow-like,
with a smaller tumid area occupying the center, medi-
al area tumid, posterior margin of each plate a thin,
membranous ridge, postlamellar ridge long, greatly
surpassing hind margin of lamina, its Y-shaped junc-
ture at distal 0.30 of sternum. Abdomen 37-40 mm.
GARRISON: Revision of Erpetogomphus
Diagnosis
Erpetogomphus boa is most similar to E. cophias, but
is easily distinguished by the thoracic pattern and the
male caudal appendages. A well-defined combined
antehumeral and humeral stripe is present in £. boa,
but no such stripe is present in E. cophias. The supe-
rior appendages of E. cophias possess a large ventral
ridge which ends in a prominent tooth 0.30 the
length of the appendage (fig. 116). This structure is
lacking in E. boa (fig. 115). Dorsally, the apical 0.30
of the superior appendages of E. cophias contains a
few strong bristles, not as many or as stout as in £.
boa.
The females of these two species are similar mor-
phologically, but may be distinguished by the thorac-
ic pattern. The postocciput of £. boa (fig. 164) does
not have the posterior depressions on each side found
in E. cophias (fig. 165).
Females of E. boa superficially resemble females of
E. viperinus, but the latter have a pair of circular de-
pressions on abdominal sternite 9 posterior to the
juncture of the postlamellar ridge which are absent in
E. boa. The vulvar lamina of £. viperinus (fig. 184) is
totally unlike that of £. boa (figs. 195-196).
Remarks
This species has been known from only two speci-
mens: Selys’ type and an incomplete male described
by Ris (1917). The lectotype is a teneral male with
the apical 0.5 of the cerci missing (fig. 114), but it
was apparently complete at the time of the original
description. The original description of the male is as
follows (translated from the French): ‘Pterostigma
light yellow brown. Head and thorax yellow, femora
yellow, with an external brown stripe, short, the ante-
rior four tarsi brownish black.
‘[male]. Superior appendages swollen at their base,
with an obtuse tooth above at the end of the swelling;
their points rounded, slightly bent inwards, hairy.
Inferior appendage divided, branching a little dis-
tance, attaining 1/2 length of the superior. Occiput
nearly straight. Tibiae brown.’
Selys’ (1859) description of the female led Calvert
(1905) to believe that it was really a specimen of £.
crotalinus, and my examination of this female con-
firms Calvert’s assertion. Erpetogomphus crotalinus ap-
pears to be restricted to the Pacific side of Mexico,
and I doubt that the locality (Vera Cruz’) is correct
for this specimen. In the supplement to the Biologia
(Calvert, 1907), Calvert received drawings of the in-
complete abdominal appendages of the type which
led him to remark that £. boa seemed hardly different
from E. elaps. Ris (1917) provided a history of the
type male of £. boa and remarked (translated from the
German): ‘I examined a long time ago in the
Hamburg Museum an old male that was different
2371
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
from elaps and corresponds with the description of
boa, that its identity [with boa] seemed likely. The lo-
cality is, unfortunately, uncertain, the label reads
‘Agua Caliente’ without further information. Its pres-
ervation is fair, the tip of the inferior appendages are
missing about the distal fourth as shown in the fig-
ure...
Examination of the hamules of the lectotype veri-
fies the identity of a series of E. boa all collected in
southern Veracruz; and the Agua Caliente specimen
of Ris is also Æ. boa.
Calvert (1905) postulated that £. sipedon might be
the female of E. boa. Although I have not seen a pair
of E. boa in copula or in tandem, I am confident in as-
cribing these females to £. boa. The similarity of the
vulvar laminae in £. boa and E. cophias (whose males
are closely related) further strengthens my conviction
that £. sipedon does not represent the female of E.
boa. Lopez (in litt.) has observed tandem pairs, con-
firming the female sex of this restricted species.
The male epiprocts, which are robust and form a
sharp point, are apparently responsible for structural
damage to the vertex of many females during mating.
Of the 26 females I examined, 14 (54%) had two
holes medially to the postoccipital tubercles. Some of
these specimens had encrusted haemolymph sur-
rounding the wounds. Dunkle (1984, 1991) reviews
cases of traumatic mating among other anisopteran
Odonata, but their negative effect, if any, is un-
known. Of the remaining 12 females, which were un-
injured, 5 were teneral and had not yet mated.
Venational details of lectotype male: fifth anteno-
dal thickened in fore wings, fourth in hind wings;
marginal cells behind fore wing paranal cells: 12/2;
anx: fore wing 14/14, hind wing 10/10; pnx: fore
wing 10/10, hind wing 10/11; cs under pterostigma:
fore wing 5/6, hind wing 5/5; anal triangular cells: 4.
Hind wing 34 mm.
Biology. — Most specimens were obtained at a city
park near Jalapa. Individuals were found by flushing
them from low vegetation about 50 m from a small
stream. Adults spent most of their time resting and
were difficult to detect among the green foliage. Its
habits are similar to those reported for E. viperinus.
Adults prefer parts of small, low volume sandy-bot-
tomed streams where banks are often covered by
trees. Reproductive maturation continues through
July. Copulation apparently occurs primarily in open
areas near streams. Ensuing pairs fly some distance
from these sites. Some males settle at the edge of the
stream, but passing females are unreceptive there.
Females, like other members of this genus, oviposit
unattended, flying over the water, stopping briefly to
drop eggs from a height of about 40 cm. López has
observed E. boa captured by spiders (Araneidae) and
robber flies (Asilidae).
238
Collection dates are July and August. Altitudes of
capture range from 1150 m (Orizaba) to 1300 m
(Jalapa).
Distribution (fig. 207). — Erpetogomphus boa has
been taken only in central Veracruz between 18°50’N
and 19°30’N along the eastern foothills of the Sierra
Madre Oriental.
Material
Type data. — Lectotype male by present designa-
tion with following data: small green label “Vera
Cruz/Salle’, yellow rectangular label: ‘ERPET. BOA / 3”
[this last label probably added by Selys during or after
- 1879], all in Selys’ hand; two pencilled labels on red
tags: ‘No. 100’; and two pencilled white labels: ‘28’,
all in an unknown hand. The caudal appendages and
accessory genitalia are shown in fig. 114. The female
specimen originally described as the female of £. boa
is actually £. crotalinus and possesses the following la-
bels: small green label: “Vera Cruz/Salle’; and white
label: ‘Herp. boa S. | 2’, both in Selys’ hand. Both
specimens in IRSN.
Other material (136, including lectotype d;
262). — Mexico: Veracruz: Parque Javier Clavijero,
Jalapa, 1300 m, 21 June 1980 (Raul López), 1d
(RWG); (same data), 11 Aug. 1980 (A. Garcés), 16
(UNAM); (same data), 13 Aug. 1980 (R. López), 1d
(UNAM); (same data), 16 Aug. 1980 (A. Garcés), 14
(UNAM); (same data), 20 Aug. 1980, 12 (UNAM);
(same data), 5 Aug. 1981, (R. López), 2d, 62 (rw);
(same data), 6 Aug. 1981, 1d (UNAM); (same data), 1
Aug. 1982, 36 , 19 (rwe); (same data), 9 Aug. 1982,
19 (UNAM); (same data), 19 Aug. 1982, 32 (UNAM);
(same data), Bosque Mesofilo de Montaña, 12 June
1981 (R. López), 19 (unam); Barranca de Cayoapa,
Teocelo, (no collector), 12 (RwG); 4.8 mi N of
Coscomatopec, (M. A. Ortiz, O. S. Flint), 16, 29
USNM); 2.5 mi S of Huatusco, 23 July 1966 (M. A.
Ortiz, O. S. Flint), 19 (usnm); 4.7 mi N of
Huatusco, Puente Ruiz Cortines, 31 July 1966 (M.
A. Ortiz, O. S. Flint), 39 (usnm); Orizaba, Ojo de
Agua, 3800 ft. (D. R. Paulson), 16, 59 (DRP, FSCA).
Erpetogomphus cophias Selys
(figs. 27-thx, 46-abd, 69-hamules, 91-penis, 116-
app, 165-vertex, 197-vl, 207-distr)
Erpetogomphus cophias Selys, 1858: 332 (72 sep.) (descr. d
‘Le Mexique, d’aprés un male du Museum de Paris’). —
Selys 1859: 537 (11 sep.) (descr. 5); Selys 1869: 175 (12
sep.) (descr. 2); Selys 1873b: 519 (75 sep.) (list); Calvert
1899: 386 (mentioned); Calvert 1905: 164 (descr. à,
2); Calvert 1907: 398 (mention of new figure); Calvert
1909: 481 (seasonal distr.); Muttkowski 1910: 86 (cat.);
Williamson and Williamson 1930: 12 (summary of stat-
us); Montgomery 1973: 239 (comment on name);
Paulson 1982: 255 (Mex.); Davies and Tobin 1985: 27
(cat.); Tsuda 1986: 87 (cat.); Bridges 1991: VII.52 (cat.);
Tsuda 1991: 95 (cat.).
Gomphus cophias. — Hagen 1861: 100 (descr. 3).
Herpetogomphus cophias. — Walsh 1862: 389 (mentioned);
Hagen 1875a: 42 (cat.); Selys 1879: 64 (2 sep.) (charac-
ters of genus); Kirby 1890: 60 (cat).
Description
Neotype male. — Entire face pale green; base of
mandibles pale green, tips black; antefrons pale green
with wash of brown at base; vertex brown, with some
green on tubercle behind each lateral ocellus; two
prominent cone-shaped pits anterolateral to median
ocellus; scape and pedicel dark brown, flagella mis-
sing; occiput broad, its dorsal surface evenly convex,
especially medially, its posterior margin slightly arcu-
ate; crest with long brown hairs; postocciput light gr-
een, evenly concave, rear of head red brown.
Prothorax primarily brown, with light green medi-
ally on anterior, median, and posterior lobes; synthor-
ax entirely pale green with wash of brown ventrally
above coxae. Coxae and trochanters pale grey green,
femora pale grey green becoming tawny distally, a
dark brown streak on lateroextensor surfaces of femo-
ra occupying almost all of profemora, about 0.60 of
mesofemora, and distal 0.30 of metafemora; tibiae,
tarsi, and armature black. Wings hyaline, venation
dark brown basally, black distally; anterior margin of
costa pale yellow to proximal end of pterostigma; pte-
rostigma brown, veins bordering it black.
Venational details. Fifth antenodal thickened in all
wings; no marginal cells behind fore wing paranal
cells; anx: fore wing 13/13, hind wing 9/9; pnx: fore
wing 7/7, hind wing 7/9; cs under pterostigma: fore
wing 5/5, hind wing 5/4; anal triangular cells: 4 .
Hind wing 29 mm.
Abdomen predominantly pale with following dark
brown markings: vestige of midlateral stripe on seg-
ment 1; this stripe more defined on segment 2, espe-
cially behind auricle; annulus; midlateral stripe on seg-
ment 3, transverse carina and darker annulus;
segments 4-6 similar to segment 3 but stripes darker,
each beginning at a little beyond beginning of seg-
ment, enlarging at transverse carina, narrowing and
then widening, especially posteriorly, but not touch-
ing dorsally; segment 7 with transverse carina and pos-
terior 0.40 of segment laterally; segment 8 pale yellow
brown with ill-defined dorsolateral dark stripe; seg-
ment 9 similar to segment 8, but with a longitudinal
lateral middorsal spot on posterior 0.75 of segment;
segment 10 pale ocher with dark red brown along an-
terior 0.20 of segment. Pale areas of segments 1-7 pri-
marily pale olive dorsally and ventrally; pale areas of
segments 8-10 red brown dorsally and ventrally.
Abdomen 35 mm.
Cercus yellow brown, linear, slightly concave dor-
sally beyond basal 0.30, tip smoothly rounded, ventral
GARRISON: Revision of Erpetogomphus
margin with inferior carina at basal 0.30, terminating
in a distinct ventral tooth; dorsal concave area of cerci
with scattered thick bristles. Epiprocts yellow brown,
distal 0.50 curved dorsally at 90°, as is typical of ge-
nus, tips slightly divergent, each with a thick, blunt
point.
Accessory genitalia. Anterior hamule black, gla-
brous, deeply forked, with posterior branch slightly
longer than anterior, the whole structure resembling a
talon; distal 0.20 of anterior branch with a longitudi-
nal, obtuse V-shaped area. Posterior hamule spatulate,
pale green, becoming brown near tip; in lateral view,
tip obtusely pointed with a small, blunt cephalad di-
rected tooth on rear margin. Peduncle of penis with
prominent rounded foliate erect lateral lobes, its pos-
terior margin not prominent, forming a gentle bilobed
area; penis with lateral lobes poorly developed posteri-
orly, forming a serrated semicircle; membranous hood
not overlapping; cornuae well developed, parallel sid-
ed, their tips evenly rounded.
Female. — Overall colouration as in male, head with
prominent conical pits anterolateral to median ocellus
as in male; vertex with tumid oval area posterior to lat-
eral ocelli; occiput narrow, crest prominent, straight to
slightly sinuate along medial 0.30, lateral arms bent
posteriorly; prominent transverse postoccipital pit
(fig. 165) immediately behind lateral arms of crest,
medial area of postocciput tumid.
Pro- and synthorax as in male, femora with brown
areas reduced with little brown on extensor surface of
metafemora.
Venational statistics (n = 2): number of marginal
cells behind fore wing paranal cells: 2-4/1-3; anx: fore
wing 13/13, hind wing 9-11/9-10; pnx: fore wing 9/8-
9, hind wing 9-10/10; cs under pterostigma: fore wing
5/4-5, hind wing 4-6/4-5. Hind wing 32-34 mm.
Abdomen with brown dorsolateral stripe more re-
duced than male, more prominent and expanded
around lateral carinae; segments 8-9 tawny, cercus
pale ocher. Vulvar lamina (fig. 197) membranous, di-
agonally corrugated; posterior margin of each plate
gently curved, a prominent ridge along its border, a
prominent long diagonal depression immediately an-
terior to posterior margin; plates meeting medially at
tumid area; cleft between plates a small notch; junc-
ture of Y-shaped postlamellar ridge well posterior to
vulvar laminar plates; area on each side of central
stem of postlamellar ridge darkened, with a slight de-
pression. Abdomen 35-37 mm.
Diagnosis
Erpetogomphus cophias is most similar to E. boa and
is diagnosed under that species. The characteristic
ventral tooth of the cercus in the male and the post-
occipital depressions in the female are autapomorphic
characters.
255)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Remarks
This is apparently a rare species. Williamson and
Williamson (1930) stated that there were only 10
known specimens. Only one other specimen from
Michoacan, collected in 1941, has come to my atten-
tion. I have examined only 3 males and 4 females, of
which one male and one female are teneral and in
poor condition. I can see no noticeable differences
among the few specimens examined.
Variation. — Venational details of one additional
male: number of marginal cells behind fore wing par-
anal cells: 3/2; anx: fore wing 11/12, hind wing 9/8;
pnx: fore wing 9/8, hind wing 9/10; cs under pteros-
tigma: fore wing 5/5, hind wing 5/5; number of anal
triangular cells 3/3. Hind wing 30 mm.
Biology. — Nothing is known of the biology of this
species.
Distribution (fig. 207). — Like £. boa, E. cophias
has a restricted distribution and apparently replaces
E. boa in the highlands of west-central Mexico south
of 20°N and west of 100°W. Records indicate an ele-
vational gradient of 1525 m (Cuernavaca) to 2438 m
(Omilteme) and flight during June and July.
Material
Type data. According to Dr. J. Legrand ( im itt. 11
May 1984), the holotype male from ‘Le Mexique,
d’après un mâle du Museum de Paris’, is missing, the-
re remaining only a large rectangular green label with
[handwritten] “E. cophias, Selys’ which was originally
attached to the specimen. Dr. Legrand was kind
enough to send a male of which he states, ‘...we have
from ex Martin’s collection a specimen called O. cop-
hias Selys (determined by Selys himself, according to
Martin...),’ which he suggested I designate as neoty-
pe. This I now do. The neotype male contains the fol-
lowing data: white label in R. Martin’s hand:
‘Ophiogomphus | cophias Selys/ Det. De Selys/
Mexique’, small green label printed: ‘MUSEUM PARIS /
Coll. R. MARTIN 1920’, and red rectangular label
handwritten: “Erpetogomphus | cophias Selys 8 / NEO-
TYPE / des./ R. W. Garrison 1986’. The abdomen is
detached and is in a triangular envelope pinned bene-
ath the specimen.
Other material (34, 49, including neotype d). —
Mexico: Guerrero: Omilteme, 8000 ft., July 1888
(H. H. Smith), 12, (BMNH); Michoacan: Tancitaro,
6000 ft. (1846 m), 4th Hoogstraal Mexican
Biological Expedition, 28 July 1941 (H. Hoogstraal),
1d (ummz); Morelos. Cuernavaca, June 1897 (O. W.
Barrett), 16, 19 (ansp); June 1888 (H. H. Smith),
16 (BMNH); 8 July 1900 (C.C. Deam), 22 (ummz).
ACKNOWLEDGEMENTS
Completion of this work would have been impossible
240
without the help of the curators of many institutions and
many individuals who placed many specimens at my dispo-
sal. I thank Donald Azuma of the Academy of Natural
Sciences, Philadelphia (Ansp); Stephen Brooks, British
Museum (Natural History) (BMNH); Dr. Paul Dessart,
Institut Royal des Sciences Naturelles de Belgique, Brussels
(IRSN); Dr. David Furth, Museum of Comparative Zoology,
Harvard University (mczc); Dr. Jean Legrand, Muséum
National d'Histoire Naturelle, Paris (MNHP); Mr. Mark
O’Brien and Mrs. Leonora Gloyd, University of Michigan,
Museum of Zoology, Ann Arbor (ummz); Dr. Oliver S.
Flint, Jr, National Museum of Natural History,
Smithsonian Institution, Washington, D. C. (usnm); Prof.
Dr. Kurt Giinther, Zoologisches Museum, Humboldt
Universität, Berlin (ZMHB), for allowing me the privilege of
examining type material in their great institutions. Dr. Jean
Belle, Velp, The Netherlands, kindly interceded on my be-
half in obtaining the syntypes of Erpetogomphus constrictor
from the Senckenberg Museum, Frankfurt-am-Main (sMF)
so that I could designate a lectotype. Examination of these
specimens allowed me to correct errors in earlier descrip-
tions and to make correct association of at least one species
with its name.
The following individuals provided specimens, records,
and other information which added considerably to this
paper: Frank Carle, Warren, NJ, for material he borrowed
from the American Museum of Natural History, NY
(AMNH); Donald Azuma, Academy of Natural Sciences,
Philadelphia (Ansp); Stephen Brooks, British Museum
(Natural History); Dr. Richard Baumann, Brigham Young
University, Provo, UT (Byuc); Dr. Paul Arnaud, California
Academy of Sciences, San Francisco (casc); Carl Cook,
Center, KY (cc); Mrs. Tineke Boomsma, Orange Walk,
Belize (TB); Jerrell Daigle, Tallahasse (jp); Dr. Alan Hardy,
California Department of Food and Agriculture,
Sacramento (cpFa); M. en C. Carlos Esquivel, San José,
Costa Rica (ce); Dr. Clifford Johnson, University of
Florida, Gainesville (cy); Dr. Boris Kondratieff, Colorado
State University, Fort Collins (csuc); Dr. James Liebherr,
Cornell University, Ithaca, NY (curc); Dr. D. A. L. Davies,
Cambridge, United Kingdom (parp); Dr. Dennis Paulson,
Burke, Washington (pre); Dr. Minter J. Westfall, Jr. and
William Mauffray (wm), Florida State Collection of
Arthropods (rsca) and International Odonata Research
Institute (tort), University of Florida, Gainesville; Dr. Jean
Belle, Velp, The Netherlands (jp) [collection now in
National Museum of Natural History, Leiden, The
Netherlands (RMNH)]; Dr. Ken Tennessen, Florence, AL
(Kr); the late Dr. Charles Hogue, Los Angeles Co. Natural
History Museum, Los Angeles (LACM); Prof. Dr. Paul
Dessart, Institut Royal des Sciences Naturelles, Brussels;
Jeffrey Cole, Los Angeles; Dr. David Furth, Museum of
Comparative Zoology, Harvard University; Dr. Michael
May, Rutgers University, NJ; Dr. Jean Legrand, Muséum
National d'Histoire Naturelle, Paris; Paul Miliotis,
Westford, MA (psm); Philippe Machet, St.-Cloud, France
(PM); Larry Muller, El Segundo, CA; Dr. Sidney Dunkle,
Collin County Community College, Plano, TX (swp); Dr.
Thomas W. Donnelly, State University of New York,
Binghamton (twp); Mark O’Brien and Leonora Gloyd,
University of Michigan, Museum of Zoology, Ann Arbor;
M. en C. Enrique González S. and Lic. Raul Armando
López P., Universidad Nacional Autónoma de México,
Mexico City (IBUNAM) and Museo Zoologica ‘Alfonso
Herrera’, Facultad Ciencias (UNAM); M. en C. Rodolfo
Novelo Gutierrez, Instituto de Ecologia, Xalapa, Veracruz,
Mexico; Dr. Oliver S. Flint, Jr., United States National
Museum, Smithsonian Institution, Washington, D. C.;
Victor Hellebuyck, Sherbrooke, Québec (vH); Prof. Dr.
Kurt Günther, Zoologisches Museum, Humboldt
Universitat, Berlin, Clark Shiffer, State College, PA; Dick
Swinney, Glendora, CA. Specimens in my collection are la-
belled rwe.
Versions of this paper benefited from critical readings by
Drs. J. Belle, T. W. Donnelly, S. W. Dunkle, M. en C.
Enrique Gonzalez S., and Lic. Rodolfo Novelo G. Lic. Raul
López P., and Lic. Rodolfo Novelo G. of Universidad
Nacional Auténoma de México very kindly made available
to me notes on the biology and reproductive behaviour of
certain Mexican Erpetogomphus Lic. Alonso Ramirez,
Universidad Nacional, Heredia, Costa Rica, supplied me
with records of E. constrictor from Costa Rica; and Dr. Jerry
Louton, United States National Museum, Smithsonian
Institution, was helpful in reviewing the cladistic analysis. I
also benefited from discussions on the philosophy of cladis-
tics from Don Frack, Covina, CA.
My wife, Jo, typed the paper and has given freely of edi-
torial advice, encouragement, and help in making time for
me to work on this project. Dr. Jan van Tol, National
Museum of Natural History, Leiden, The Netherlands
(RMNH) provided editorial advice and supplemented my
cladistic analysis and interpretation as this paper was going
to press. To all these persons, I express my sincere thanks
and appreciation for all the help they have rendered me.
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de Belgique (2) 28: 168-208 (1-45 separate).
Selys-Longchamps, Edm. de, 1873a. Troisièmes additions
au synopsis des Gomphines. — Bulletin de l'Académie
royale de Belgique (2) 35: 732-774 (1-46 separate).
Selys-Longchamps, Edm. de, 1873b. Appendices aux
troisièmes additions et liste des Gomphines, descrites
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l’Académie royale de Belgique (2) 36: 492-531 (47-87
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Selys-Longchamps, Edm. de, 1878. Quatrièmes additions
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Selys-Longchamps, Edm. de, 1879. Revision des
Ophiogomphus et descriptions de quatre nouvelles
GARRISON: Revision of Erpetogomphus
Gomphines Americaines. — Comptes Rendus de la Socié-
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245
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Mesa, Arizona (Odon.). Entomological News 25: 225-
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Received: 19 November 1993
Accepted: 10 August 1994
246
GARRISON: Revision of Erpetogomphus
1. constrictor d
9. eutainia d
7. eutainia d
10. leptophis d
13. elaps 3 (TA Niopeltisnd
Figs. 1-14. Synthorax of Erpetogomphus species, lateral view (males, except given otherwise). — 1, constrictor (Honduras); 2, sa-
baleticus (holotype); 3, tristani (Costa Rica); 4, ophibolus (Mexico: Veracruz State); 5, agkistrodon (female holotype); 6, schau-
si (holotype); 7, eutainia (Texas: Gonzales Co.); 8, eutainia (Mexico: Veracruz State); 9, eutainia (Costa Rica); 10, leptophis
(holotype); 11, elaphe (holotype); 12, elaps (Mexico: Sinaloa State); 13, elaps (Mexico: Morelos State); 14, bopeltis (holotype).
Scale line: 3 mm.
247
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
4
we
23. heterodon d
26. boa 3
SSE 27. cophias
Figs. 15-27. Synthorax of Erpetogomphus species, lateral view (males, except given otherwise). — 15, bothrops (holotype); 16,
viperinus (Mexico, Veracruz State); 17, designatus (lectotype); 18, designatus (Mexico, Durango State); 19, sipedon (female ho-
lotype); 20, / lampropeltis (holotype); 21, lampropeltis natrix (holotype); 22, crotalinus (lectotype); 23, heterodon (holotype);
24, compositus (California, Yolo Co.); 25, compositus (paratype of coluber); 26, boa (Mexico: Veracruz State); 27, cophias
(Mexico: Morelos State). Scale line 3 mm.
248
GARRISON: Revision of Erpetogomphus
28. constrictor 9
29. ophibolus 3
30. agkistrodon ?
31. schausi d
32. eutainia d
33. eutainia 9
34. leptophis d
35. leptophis ©
1 i z I EHEN;
36. liopeltis d di iby SENSI =;
JSS
Figs. 28-36. Abdomen, lateral view. — 28, constrictor £ (Honduras); 29, ophibolus $ (Mexico: Veracruz State); 30, agkistro-
don 2 (holotype); 31, schausi 3 (holotype); 32, ewtainia 3 (Texas: Gonzales Co); 33, eutainia ® (idem); 34, leptophis 3 (ho-
lotype); 35, leptophis 2 (allotype); 36, liopeltis 3 (holotype). Scale line 10 mm.
249
Sie
38.
39}
40.
41.
42.
43.
44.
45.
46. cophias d
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
= à
bothrops d
viperinus d
sipedon d
sipedon ©
crotalinus d
heterodon d
heterodon 2
boa 6
boa 2
Figs. 37-46. Abdomen, lateral view. — 37, bothrops 3 (holotype); 38, viperinus d (Mexico: Veracruz State); 39, sipedon 4
(Mexico: Morelos State); 40, sipedon 2 (holotype); 41, crotalinus & (Mexico: Jalisco State); 42, heterodon 3 (holotype); 43,
heterodon 2 (allotype); 44, boa & (Mexico: Veracruz State); 45, boa ® (idem); 46, cophias & (Mexico: Morelos State). Scale
line 10 mm.
250
GARRISON: Revision of Erpetogomphus
47. constrictor 48. sabaleticus
52. eutainia 53. leptophis
. elaps
oz ET
57. liopeltis 58. bothrops
Figs. 47-58. Anterior and posterior hamules. First view is anterolateral view showing profile of anterior hamule; second view
(when shown) is lateral view showing profile of posterior hamule. — 47, constrictor (Mexico: San Luis Potosi State); 48, saba-
leticus (holotype); 49, tristani (Costa Rica); 50, ophibolus (Mexico: Veracruz State); 51, schausi (holotype); 52, eutainia (Texas:
Gonzales Co.); 53, leptophis (holotype); 54, elaphe (Costa Rica); 55, elaps (holotype); 56, elaps (variant) (Mexico: Chiapas
State); 57, liopeltis (paratype); 58, bothrops (paratype). Scale line 2 mm.
Do
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
y
61. sipedon
62. lampropeltis 63. lampropeltis natrix 64. crotalinus
lampropeltis
68. boa 69. cophias
Figs. 59-69. Anterior and posterior hamules. First view is anterolateral view showing profile of anterior hamule; second view
(when shown) is lateral view showing profile of posterior hamule. — 59, viperinus (Mexico: Veracruz State); 60, designatus (lec-
totype); 61, sipedon (Mexico: Morelos State); 62, / lampropeltis (California: Ventura Co.); 63, / natrix (holotype); 64, crot-
alinus (Mexico: Michoacan State); 65, heterodon (New Mexico: Catron Co.); 66, compositus (Arizona: Pima Co.); 67, compo-
situs (paratype of coluber); 68, boa (lectotype); 69, cophias (Mexico: Morelos State). Scale line 2 mm.
DD
GARRISON: Revision of Erpetogomphus
EN
74. schausi
N
79. liopeltis
NS
80. bothrops
Figs. 70-80. Penis segment 4, left: dorsal view, right: lateral view. — 70, constrictor (Mexico: San Luis Potosi State); 71, saba-
leticus (holotype); 72, tristani (Costa Rica); 73, ophibolus (Mexico: Veracruz State); 74, schausi (holotype); 75, eutainia (Texas:
Gonzales Co.); 76, leptophis (holotype); 77, elaphe (Costa Rica); 78, elaps (holotype); 79, liopeltis (paratype); 80, bothrops
(Mexico: Tamaulipas State). Scale line 1 mm.
253
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
85. lampropeltis natrix KL 86. crotalinus
j
91. cophias
Figs. 81-91. Penis segment 4, left: dorsal view, right: lateral view. — 81, viperinus (Mexico: Veracruz State); 82, designatus
(Texas: Gonzales Co.); 83, sipedon (Mexico: Morelos State); 84, / lampropeltis (California: Ventura Co.); 85, / natrix
(Arizona: Cochise Co.); 86, crotalinus (Mexico: Michoacan State); 87, heterodon (New Mexico: Catron Co.); 88, compositus
(CA: Yolo Co.); 89, compositus (paratype of coluber); 90, boa (lectotype); 91, cophias (Mexico: Morelos State). Scale line 1
mm.
254
GARRISON: Revision of Erpetogomphus
92. constrictor 93. sabaleticus 94. tristani 95. ophibolus
96. schausi 97. eutainia 98. leptophis 99. elaphe
a 101. liopeltis 102. bothrops 103. viperinus
Figs. 92-103. Caudal appendages of male, above = lateral view, below = dorsal view of cerci. — 92, constrictor (Mexico:
Tamaulipas State); 93, sabaleticus (holotype); 94, tristani (holotype); 95, ophibolus (Mexico: Veracruz State); 96, schausi (ho-
lotype); 97, eutainia (Texas: Gonzales Co.); 98, leptophis (holotype); 99, elaphe (Costa Rica); 100, elaps (holotype); 101, Lo-
peltis (holotype); 102, bothrops (holotype); 103, viperinus (lectotype). Scale line 5 mm.
255
“TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
106. sipedon
n ? Se 2
107. lampropeltis
lampropeltis
104. designatus
110. heterodon
108. lampropeltis
natrix
A ee =
112. compositus 113. compositus 116. cophias
i =) (3 | |
117. liopeltis 118. bothrops 119. viperinus 115. boa
Fig. 104-116. Caudal appendages of male, above = lateral view, below = dorsal view of cerci. — 104, designatus (lectotype);
105, sipedon (Mexico: Morelos State); 106, sipedon (only lateral view) (Mexico: Durango State); 107, / lampropeltis
(California: Ventura Co.); 108, Z natrix (holotype); 109, crotalinus (lectotype); 110, heterodon (holotype); 111, heterodon
(Mexico: Durango State); 112, compositus (California: Yolo Co.); 113, compositus (paratype of coluber); 114, boa (only lateral
view) (lectotype); 115, boa (Mexico: Veracruz State); 116, cophias (Mexico: Morelos State). Figs. 117-119. Right cercus, me-
diolateral view. — 117, liopeltis (Mexico: Nuevo Leon State); 118, bothrops (Mexico: Nayarit State); 119, viperinus (Mexico:
Veracruz State). Scale line 5 mm.
256
GARRISON: Revision of Erpetogomphus
( aX
120. tristani 121. ophibolus 122. eutainia 123. eutainia
124. eutainia 125. eutainia 126. liopeltis
127. bothrops 128. bothrops 129. viperinus
130. designatus 131. designatus 132. lampropeltis
lampropeltis
133. crotalinus 134. compositus 135. boa
Figs. 120-135. Erpetogomphus. — 120, tip of epiproct of male, posterior view, tristani (Costa Rica); 121, idem, ophibolus
(Mexico: Veracruz State); 122, epiproct, lateral view, eutainia (Texas: Gonzales Co.); 123, idem, dorsolateral view; 124-135,
tip of epiprocts of male, posterior view, 124, eutainia (Texas: Gonzales Co.); 125, eutainia (Guatemala); 126, Liopeltis
(Mexico: Nuevo Leon State); 127, bothrops (Mexico: Nayarit State); 128, bothrops (holotype); 129, viperinus (Mexico:
Veracruz State); 130, designatus (Texas: Gonzales Co.); 131, designatus (Mexico: Durango State); 132, / lampropeltis
(California: Ventura Co.); 133, crotalinus (Mexico: Jalisco State); 134, compositus (Arizona: Coconino Co.); 135, boa
(Mexico: Veracruz State). Scale line 2 mm (figs. 122-123), 1 mm (figs. 120-121, 124-135).
Dj
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
SO
136. bothrops
DÌ 0
138. lampropeltis
lampropeltis d
139. compositus d 141. liopeltis
145. constrictor 146. tristani 147. ophibolus
TA
N pi
153. liopeltis
151. leptophis “452. elaps |
Figs. 136-153. Erpetogomphus. — 136-139. Anterior part of frons, dorsal view. 136, bothrops 2 (Mexico: San Luis Potosi
State); 137, viperinus 2 (Mexico: Veracruz State); 138, /. lampropeltis 3 (California: Ventura Co.); 139, compositus 3 (AZ:
Maricopa Co.). — Figs. 140-141. Crest outline of occiput, dorsal view. 140, liopeltis (holotype), 141, /opeltis (Mexico: Nuevo
Leon State). — Figs. 142-144. Head, anterior view. 142, schausi (holotype); 143, eutainia (Costa Rica); 144, eutainia (Texas:
Gonzales Co.). — Figs. 145-153.Vertex, occiput of female, dorsal view. 145, constrictor (Honduras); 146, tristani (Costa Rica);
147, ophibolus (Mexico: Veracruz State); 148, agkistrodon (holotype); 149, schausi (Costa Rica); 150, eutainia (Texas:
Gonzales Co); 151, leptophis (allotype); 152, elaps (Mexico: Morelos State); 153, opeltis (allotype). Scale line 4 mm (figs.
142-144); 1 mm (136-141, 145-153).
258
GARRISON: Revision of Erpetogomphus
157. designatus 158. sipedon 156. designatus
LV
159. sipedon
160. lampropeltis 161. crotalinus
lampropeltis
166. viperinus
OAS KAREN ni
165. cophias
Figs. 154-165. Vertex, occiput of female, dorsal view. 154, bothrops (allotype); 155, viperinus (Mexico: Veracruz State); 156,
designatus (paralectotype); 157, designatus (Mexico: Durango State); 158, sipedon (form ‘b’) (Mexico: Puebla State); 159, si-
pedon (holotype); 160, £ lampropeltis (California: Ventura Co.); 161, crotalinus (paralectotype); 162, heterodon (allotype);
163, compositus (holotype); 164, boa (Mexico: Veracruz State); 165, cophias (Mexico: Guerrero State). — Figs. 166-167. Crest
and part of rear of head (postocciput) of female, dorsal view. 166, viperinus (Mexico: Veracruz State); 167, boa (Mexico:
Veracruz State). Scale line 1 mm.
259
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
sd
Yr viv dy!
171. constrictor 172. sabaleticus 173. tristani 174. tristani
175. ophibolus 176. agkistrodon 177. schausi 178. eutainia
179. leptophis 180. elaphe 181. elaps 182. liopeltis
Fig. 168-170. Metathoracic leg, lateral view. — 168, sipedon 2 (holotype); 169, crotalinus 3 (lectotype); 170, heterodon à
(holotype). — Figs. 171-182. Vulvar lamina, dorsal view. — 171, constrictor (Honduras); 172, sabaleticus (allotype); 173, tri-
stani (allotype); 174, tristani (Costa Rica); 175, ophibolus (Mexico: Veracruz State); 176, agkistrodon (holotype); 177, schausi
(Costa Rica) (distorted on right side); 178, eutainia (Mexico: Veracruz State); 179, leptophis (allotype); 180, elaphe (Costa
ae? 181, elaps (Mexico: Morelos State); 182, liopeltis (Mexico: Nuevo Leon State). Scale line 4 mm (figs. 168-170), 2 mm
(figs. 171-182).
260
GARRISON: Revision of Erpetogomphus
183. bothrops 184. viperinus 186. designatus
187. sipedon 188. sipedon 189. lampropeltis 190. lampropeltis
lampropeltis natrix
191. crotalinus 193. heterodon
A
N
i A ST) :
195. boa 196. boa 197. cophias
Figs. 183-197. Vulvar lamina, dorsal view. — 183, bothrops (allotype); 184, viperinus (Mexico: Veracruz State); 185, designa-
tus (paralectotype); 186, designatus (Texas: Gonzales Co.); 187, sipedon (holotype); 188, sipedon (paratype); 189, 1 lampro-
peltis (California: Ventura Co.); 190, / natrix (allotype); 191, crotalinus (paralectotype); 192, crotalinus (Mexico: Jalisco
State); 193, heterodon (allotype); 194, compositus (Arizona: Maricopa Co.); 195, boa (Mexico: Veracruz State); 196, boa
(Veracruz State); 197, cophias (Mexico: Guerrero State). Scale line 2 mm.
261
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
I N: È
Nez DS si EL
@ = constrictor => CS (CS x
A= ophibolus ES N
@ = eutainia
À = leptophis
M = liopeltis
@ - bothrops = |
ÀA=viperinus ~~ >
ers
B = narrow hamular
form of elaps
A- elaphe
id
i OW
SAG SEGA
| Bee BV ae
|
202
Figs. 198-202. Distribution patterns. — 198, E. ophibolus group; 199, E. eutainia group; 200, E. elaps group (in part); 201, E.
elaps group (in part); 202, E. designatus.
262
GARRISON: Revision of Erpetogomphus
© - 1. lampropeltis
A =|. natrix
© = compositus
À = heterodon
|
Figs. 203-207. Distribution patterns. 203, E. sipedon; 204, E. lampropeltis, 205, E. crotalinus, 206, E. heterodon and E. com-
positus, 207, E. boa and E. cophias.
263
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
O. severus constr. eut. elaps elaphe lampro. boa
elt hee
a
208 aosta 0
O. severus constr. eut. elaps cophias
209
O. severus
211-a i
Ophiogomphus 211-b 211-c 211-d 211-e
severus tristani viperinus compositus boa
Figs. 208-211. — 208, morphocline of anterior hamule, character states 0-5; 209, morphocline of posterior hamule, character
states 0-4; 210, morphocline of tip of posterior hamule, character states 0-5; 211, dorsal view of fourth segment of penis (S4);
figs. 211a and 211b have 4th segment about as long as wide, figs. 21 1c-e about twice as long as wide. Scale line 2 mm (figs.
208-209); 1 mm (figs. 210-212).
264
GARRISON: Revision of Erpetogomphus
Ophiogomphus ophibolus eutainia crotalinus
severus group group group
> (C i | > prepuce lacking
A (state 3)
8 prepuce
prepuce prepuce (hidden by lateral lobe:
212 (state 0) (state 1) state 2)
crotalinus
Ophiogomphus ophibolus eutainia elaps subgroup
severus group group group designatus (except
designatus)
rer: 24 T ap Fi Yale a
213 0 1 2 5 4 5
€ NX boa, cophias
ophibolus compositus \\ |
group (part) È iy)
| BE nn
i crotalinus {| | x 60
(2) EE) \ N
x lampropeltis = a E
schausi a na
= ene) % i
elaps | \ | 40
In! subgroup |
et \
eutainia ii \
II Ai
Oph. sev. <50 A
o ed i ea =
N + 2. ‘ .
eS 4
ec 7
00
214 oo PR ne meo
Ophiogomphus severus eutainia viperinus
ZI (state 0) (state 1) (state 2)
Figs. 212-215. — 212, morphocline of prepuce of penis, character states 0-3; 213, morphocline of shape of lateral lobe, char-
acter states 0-5; 214, morphocline of cornua: character states 0-6 for character 19 and character states 0-2 for character 20;
215, morphocline of spermatheca in dorsal (internal) view character states 0-2 (figures show dorsal (internal) view of sternum
9 (including postlamellar ridge). Spematheca is dorsad of vulvar lamina). Scale line 1 mm (figs. 212, 215); 0.5 mm (figs. 213-
214).
265
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
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266
GARRISON: Revision of Erpetogomphus
0 species
© 1 species
ER 2 species
«000»
3 species
4 species
5 species
8 species
Fig. 217. geographic distribution of Erpetogomphus collected in the U. S. A., Mexico, and Central America. Each square is
approximately 150 kilometers square.
267
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
=
|
j
= ophibolus/eutainia groups
| = elaps/crotalinus groups
[a = ophibolus group
Fe. soe
I ' = eutainia group
agkistrodon
ophibolus
schausi
fi
sabaleticus }
leptophis
|
|
|
223
224
Figs. 218-224. — 218, Generalized track for genus Erpetogomphus 219, generalized tracks for Erpetogomphus ophibolus | eu-
tainia and E. elaps | crotalinus groups; 220, generalized tracks for Erpetogomphus ophibolus and E. eutainia groups; 221, tracks
of species within the Erpetogomphus ophibolus group; 222, tracks of species within the Erpetogomphus eutainia group; 223, ge-
neralized tracks for Erpetogomphus crotalinus and E. elaps groups; 224, tracks of species within the Erpetogomphus elaps group
(
in part).
268
GARRISON: Revision of Erpetogomphus
iN liopeltis
226
CT SS >
LLS
Aes
229. viperinus
230. heterodon
Figs. 225-230. — 225, tracks of species within the Erpetogomphus elaps group (in part); 226, tracks of species within the
Erpetogomphus crotalinus group; 227, pair of wings of opeltis & (Mexico: Nuevo Leon State); 228, idem, bothrops 3 (Mexico:
Nayarit State); 229, idem, viperinus & (Mexico: Veracruz State); 230, idem, heterodon 3 (N. Mexico: Grant Co.)
269
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
270
HERMAN DE JONG
Institute for Systematics and Population Biology, Amsterdam
ab eiWLOGENY OF THE SUBGENUS TIPULA
(SAVTSHENKTA) (DIPTERA: TIPULIDAE), WITH
SLECIAD REFERENCE TO THE WESTERN
MEDITERRANEAN FAUNA
In memory of E. N. Savchenko (12.VII.1909 - 8.1.1994)
Jong, H. de, 1994. The phylogeny of the subgenus Tipula (Savtshenkia) (Diptera: Tipulidae),
with special reference to the western Mediterranean fauna. — Tijdschrift voor Entomologie 137:
271-323, figs. 1-202, tabs. 1-5, appendices A-C. [rssn 0040-7496]. Published 15 December
1994.
The phylogeny of the currently recognized 87 species and subspecies of the subgenus Tipula
(Savtshenkia Alexander) (Diptera, Tipulidae) is discussed in reference to 87 morphological
characters of the adults. The distribution of the species and subspecies of Savtshenkia is briefly
outlined. Taxonomic novelties include the description of three new western Palaearctic species,
viz. alpha, omega, and trinacria; the establishment of the synonymy of broweri Alexander under
fragilina Alexander, insignifica Alexander under invenusta Riedel, nebulipennis Alexander under
alpium Bergroth, packardi Alexander under invenusta Riedel, and perparvula Alexander under
ignobilis Loew; the elevation of the subspecies subsignata haennii Dufour to species rank; and
the removal of the species convexifrons Holmgren from Savtshenkia.
H. de Jong, Department of Entomology, Institute for Systematics and Population Biology
(Zoological Museum), Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands.
Key words. — Diptera, Tipulidae, Savtshenkia, phylogeny, western Mediterranean.
As part of a research project on the historical bio-
geography of the western Mediterranean region, the
phylogeny of the subgenus Tipula (Savtshenkia
Alexander) is discussed. Savtshenkia as defined here,
contains 87 described species and subspecies, of
which 22 have a restricted distribution in the western
Mediterranean. Special emphasis is laid on the estab-
lishment of the phylogenetic position of these western
Mediterranean endemics, as they offer potential clues
to a better understanding of the historical biogeogra-
phy of the area.
The species constituting the subgenus Savtshenkia
were formerly grouped in the marmorata species
group of the genus Tipula Linnaeus (Alexander
1919a, 1919b et seqq.). The same assemblage of spe-
cies has also been known as the fragilis group
(Alexander 1934 et seqq.) and obsoleta group
(Lackschewitz 1936). Usually, these species groups
were referred to the subgenus Oreomyza Pokorny
(Alexander 1934, Edwards 1931). Savchenko (1961),
misinterpreting Riedel (1913) regarding the type spe-
cies of Pterelachisus Rondani, synonymized Oreomyza
under the latter taxon. Savchenko’s lapsus resulted in
the improper use of the name Pterelachisus for the
species at present placed in Savtshenkia. Although
Savchenko’s reasoning was fallacious, Mannheims
(1962a) showed that the synonymy of Oreomyza and
Pterelachisus sensu Rondani in itself was valid and in-
troduced the name Savtshenkia to replace
Pterelachisus sensu Savchenko. As Mannheims omit-
ted to denote the type species of this new taxon,
Savtshenkia remained a nomen nudum until
Alexander (1965a) designated rufina Meigen as the
type species. (In the literature, a paper by Alexander
published in the 1965 volume of the Philippine
Journal of Science is usually cited as the relevant ref-
erence for the authorship of the name Savtshenkia. As
the pertaining publication was actually issued August
9, 1966, Alexander’s paper that appeared in Pacific
Insects of June 20, 1965, should be accepted as the
correct reference for the type designation.)
Savtshenkia has an essentially Holarctic distribu-
tion (fig. 1), with 65 species and subspecies occurring
in the Palaearctic Region and eight in the Nearctic
Region. Both regions have two species in common. In
addition to the Holarctic members of the subgenus,
five Oriental and 12 Afrotropical species and subspe-
cies are known. The Oriental fauna of Savtshenkia has
a single species in common with the Palaearctic fau-
na, the Afrotropical fauna shows no species overlap
271
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
A
Fig. 1. Global distribution of the subgenus Tipula (Savtshenkia).
with that of any other biogeographical region. At the
end of this paper a brief account of the distribution of
the species of Savtshenkia is provided. | will present
more detailed information on the distribution of the
western Mediterranean species and their closest rela-
tives in a forthcoming paper on the historical bio-
geography of the western Mediterranean. In
Appendix A, three western Palaearctic species of
Savtshenkia are described as new, five synonymies are
established, while one subspecies is raised to species
rank and one species is removed from the subgenus.
The resulting list of species and subspecies of
Savtshenkia as recognized in this paper is given in ta-
ble 1.
Savchenko (1961, under Pterelachisus and
Theowald (1957, under Oreomyza, 1967, 1973,
1978) arranged the (western) Palaearctic members of
Savtshenkia in a number of species groups.
Theowald’s 1957 and 1967 arrangements were based
on characters of the preadult stages of western
Palaearctic species and contained four species groups
(table 2). Savchenko (1961) recognized five species
groups plus a miscellaneous group within the fauna of
the former USSR (table 3). Theowald’s 1973/1978
classification of the adults of Savtshenkia in 14 species
groups basically is a refinement of Savchenko’s 1961
concept (table 3). The arrangements of both authors
will be compared with the results of the present study
in the ‘Discussion of adopted phylogeny’, below.
DD
MATERIAL, METHODS AND TERMINOLOGY
During the course of this investigation material re-
presenting 78 species and subspecies of Savtshenkia
was examined (table 1). Most specimens used for the
present study originated from the Institute for
Systematics and Population Biology (Zoological
Museum), Amsterdam. Supplementary material was
studied during a short stay at the United States
National Museum of Natural History, Smithsonian
Institution, Washington D.C. In addition to this,
material was borrowed from the following institu-
tions: Academy of Natural Sciences, Philadelphia;
Musée d'Histoire Naturelle, Neuchâtel; Natural
History Museum, London; Staatliches Museum für
Naturkunde, Stuttgart; United States National
Museum of Natural History, Smithsonian
Institution, Washington D.C.; Zoologiska
Institutionen, Systematiska avdelningen, Lund.
The majority of specimens consisted of pinned ma-
terial; in addition to this, material preserved in alco-
hol was studied from the collections in Amsterdam,
Lund, and Neuchâtel.
Preparations of the male and female terminalia
were made by removing these parts and clearing them
in a nearly boiling 10% KOH solution for about five
minutes. After rinsing with water and 70% alcohol,
the terminalia were transferred to glycerol.
Examination of the specimens and their terminalia
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
Table 1. List of species and subspecies of Tipula (Savtshenkia) as recognized in this paper. Abbreviations: +: examined; -: not
examined; u: unknown; Afro: Afrotropical Region; Nea: Nearctic Region; Or: Oriental Region; Pal: Palaearctic Region; Th.,
Duf. & Oost.: Theowald, Dufour & Oosterbroek.
3d 2 Region
aberdareica Alexander, 1956 ERLE
a. ulugurica Alexander, 1962 U ELO
akeleyi Alexander, 1956 + uly AfL
alpha sp. n. + + Pal
alpium Bergroth, 1888 + + Nea, Pal
asbolodes Speiser, 1909 + - Afro
aspromontensis Theowald, 1973 + + Pal
aster Theischinger, 1983 UTP al
atlas Pierre, 1924 + + Pal
baltistanica Alexander, 1936 + + Or, Pal
benesignata Mannheims, 1954 + eal
boreosignata Tjeder, 1969 SALE bal
breviantennata Lackschewitz, 1933 + eal
caligo Alexander, 1956 + + Afro
cheethami Edwards, 1924 + + Pal
chrysocephala Mannheims, 1958 + u Afro
confusa Van der Wulp, 1887 Dal
corsosignata Th., Duf. & Oost. 1982 + + Pal
cyrnosardensis Th., Duf. & Oost. 1982 + ger Dal
draconis Alexander, 1964 + u Afro
eleonorae Theischinger, 1978 + u Pal
elgonensis Alexander, 1956 TATTO,
eugeni Theowald, 1973 + + Pal
fragilina Alexander, 1919 + + Nea
fragilis Loew, 1863 + + Nea
gimmerthali Lackschewitz, 1925 + + Pal
g. mattheyi Theowald & Dufour, 1983 - u Pal
g. pteromaculata Th., Duf. & Oost. 1982 + u Pal
glaucocinerea Lundstrôm, 1915 a Yo) Pall
goriziensis Strobl, 1893 + + Pal
graciae Alexander, 1947 + u Nea
grisescens Zetterstedt, 1851 + Pal
haennii Dufour, 1991 stat.n. ET
hancocki Alexander, 1956 + + Afro
hartigiana Th., Duf. & Oost. 1982 RE Rll
holoptera Edwards, 1939 + + Pal
ignobilis Loew, 1863 + + Nea
imperfecta Riedel, 1914 = - Afro
interserta Riedel, 1913 + + Pal
invenusta Riedel, 1919 + - Nea, Pal
i. microinvenusta Dufour, 1990 i Ja
i. subinvenusta Slipka, 1950 + + Pal
jeekeli Mannheims & Theowald, 1959 + + Pal
kiushiuensis Alexander, 1925 ge 4 Pall
koreana Alexander, 1934 mae Pal
was carried out using a Wild stereomicroscope with a
magnification of up to 100 times. Drawings were
made with the aid of a drawing tube attached to the
microscope. Illustrations of the genital structures
were made from macerated specimens. For perma-
nent storage, the terminalia were transferred to a mi-
crovial containing a drop of glycerol. The microvial
was pinned with the relevant specimen.
The parsimony program HENNIG86, version 1.5
(Farris 1988) was used to analyse the phylogeny.
d 2 Region
letifera Alexander, 1951 = u Pal
limbata Zetterstedt, 1838 & = Pal
lundbladi Mannheims, 1962 HN Pal
macaronesica Savchenko, 1961 + u Pal
mannheimsi Theowald, 1973 sw Pal
minuscula Savchenko, 1971 2 = Pal
mohriana Alexander, 1954 + + Pal
multipicta Becker, 1908 + + Pall
nephrotomoides Alexander, 1924 + + Afro
n. invariegata Alexander, 1956 u + Afro
nielseni Mannheims, & Theowald, 1959+ + Pal
nivalis Savchenko, 1961 > =) Pall
obsoleta Meigen, 1818 gs al
odontostyla Savchenko, 1961 mo Pal
omega sp. n. + u Pal
ornata Theowald & Oosterbroek, 1987 + + Pal
pagana Meigen, 1818 + + Pal
pechlaneri Mannheims &
Theowald, 1959 + + Pal
persignata Alexander, 1945 + u Pal
p. tofina Alexander, 1945 + wu Pal
phoroctenia Alexander, 1919 + + Nea
postposita Riedel, 1919 + PI 2:11
productella Alexander, 1928 = u Nea
rufina Meigen, 1818 db db JRall
r. maderensis Lackschewitz, 1936 fo sh Pal
sardosignata Mannheims &
Theowald, 1959 Dall
sciadoptera Alexander, 1964 + u Or
serrulifera Alexander, 1942 + + Pal
signata Staeger, 1840 + + Pal
simulans Savchenko, 1966 + + Pal
sordidipes Alexander, 1961 + + Or
staegeri Nielsen, 1922 + + Pal
subalpium Savchenko, 1961 = u Pal
subnodicornis Zetterstedt, 1838 + + Pal
subsignata Lackschewitz, 1933 + +) Pal
s. cazorla Dufour, 1991 om Pal
subvafra Lackschewitz, 1936 + + Pal
tetragramma Edwards, 1928 = u Or
trinacria sp. n. + + Pal
tulipa Dufour, 1983 al
venerabilis Alexander, 1936 + u Or
villeneuvii Strobl, 1909 + + Pal
More information on the calculation procedures fol-
lowed is given in “Discussion of adopted phylogeny’.
The terms for the structures of Savtshenkia as used
in the character discussion are in general in accor-
dance with the terms employed by McAlpine (1981),
with a few additions for particular features of the
Tipulidae of the study group. The terms are ex-
plained in figs. 3-6, 12, and 58. It should be noted
that I consider the structure lateral of the gonapoph-
ysis in Savtshenkia homologous with the fragmentum
273
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
grisescens
breviantennata
tulipa
goriziensis
cheethami
simulans
ignobilis
subnodicornis
baltistanica
glaucocinerea
postposita
lundbladi
multipicta
——— Lunatipula
Mediotipula
atlas
a
FeS
œ
a
————_—— hartiglana
a
a
co is
Du ©
DO villeneuvil
w
w
= u
No ia
©
rufina rufina
rufina maderensis
nephrotomoides
elgonensis
draconis
aberdareica
asbolodes
hancocki
akeleyi
chrysocephala
macaronesica
Jeekeli
obsoleta
holoptera
pagana
7 16,58
721 J35 84,87
©
48,49
55,57
ces
x4 (eee +86
+ 53,83
Fig. 2. Strict consensus tree of 14 equally parsimonious trees derived after successive weighting for the species and subspecies
of Tipula (Savtshenkia). —: synapomorphy;
in the sense of Rees & Ferris (1939) and Frommer
(1963) and name it accordingly (figs. 3, 4).
Mannheims (1951) denoted the same sclerite A9s
(appendage of sternite 9). Suture x is employed in the
sense of Neumann (1958). A recent overview of the
morphology of the terminalia of Tipulidae was given
by Tangelder (1985) to which paper the reader is re-
ferred to for further details.
Acronyms used throughout the text are the follow-
ing:
BMNH: Natural History Museum, London,
England; BsnH: Boston Society of Natural History,
Boston, Massachusetts, U.S.A; cncı: Canadian
National Collection, Biosystematics Research Centre,
Ottawa, Ontario, Canada; mczc: Museum of
Comparative Zoology, Harvard University,
Cambridge, Massachusetts, U.S.A.; MNHN: Muséum
National d'Histoire Naturelle, Paris, France; MRAC:
Koninklijk Museum voor Midden Afrika, Tervuren,
Belgium; usNM: United States National Museum,
Smithsonian Institution, Washington D.C., U.S.A;
zıas: Zoological Institute, Russian Academy of
Sciences, St. Petersburg, Russia; ZMAN: Institute for
Systematics and Population Biology, Department of
274
: homoplasy; X: reversal.
Entomology, Zoological Museum, Amsterdam, the
Netherlands.
BIOLOGY
In general, the members of Savishenkia are adapted
to either mountainous, high latitudinal, or autumnal
conditions. Within these limits, adults of the respec-
tive species of Savtshenkia show a wide variety of pre-
ferred biotopes. They can be found in alpine
Rhododendron assocations, the heath zone at about
3300 m, mountainous wet or dry Pinus forests and
mixed woods, low woods, low moorlands, marshes,
bogs, and gardens. The species may occur in the ve-
getation near the sources and on the banks of streams,
on boulders in streams, on dripping rocks, and in the
splash zone of cascades (Alexander 1961, Audcent
1932, Brunhes & Dufour 1992, Bryce 1956,
Coulson 1959, 1962, Dufour 1983, 1986, 1990,
1991, Mannheims & Pechlaner 1963, Stubbs 1992,
Theowald 1973, Theowald, Dufour & Oosterbroek
1982).
The larvae of the species of Savtshenkia are associat-
ed with mosses and liverworts. They are recorded
immerthali pteromaculata
=
£
©
9
E
E
ao
Fi
£
=
3
E
E
immerthali mattheyi
confusa
interserta
ornata
graciae
sordidipes
venerabilis
eugeni
serrulifera
aspromontensis
aster
phoroctenia
odontostyla
fragilina
limbata
subvafra
pechlaneri
O fragilis
26
nvenusta subinvenusta
H. DE Jone: Phylogeny of Tipula (Savtshenkia)
nvenusta microinvenusta
nvenusta invenusta
cyrnosardensis
staegeri
nielseni
sardosignata
haennii
corsosignata
subsignata subsignata
subsignata cazorla
eleonorae
benesignata
persignata persignata
persignata tofina
signata
trinacria
mannheimsi
omega
9
> boreosignata
F 67
[es en
.
71
177,78
+28,29,76
30,81+14,34 72
8
+13,18,63,64
026
+62
from mosses on stones and logs in and along cold
streams, from wet moorland mosses and mosses of
cold bogs of tundra and high mountains, as well as
from relatively dry mosses on walls, stones and trees
(Gelhaus 1986).
As in most Tipulidae, the majority of species of
Savtshenkia have a one-year life cycle. A few are, or
appear to be, bivoltine (viz. alpium, breviantennata,
rufina rufina, and rufina maderensis, Dufour 1986,
Mannheims 1951, Theowald 1973). On the other
hand, Lantzov (1982) suggested a long term larval de-
velopment for the arctic species glaucocinerea of up to
six years. [Multi-annual life cycles are known of other
species of Tipulidae with a northerly distribution; see
Pritchard (1983) for an overview of life cycles in
Tipulidae]
Details on the copulation of a few species of
Savtshenkia were given by Hemmingsen (1954, 1962;
viz. limbata, signata, staegeri, and subsignata).
Hemmingsen (1952) provided a short account on
oviposition of the species confusa (as marmorata) and
rufina.
SYSTEMATIC POSITION
The species and subspecies of Savtshenkia can be
distinguished from other Tipulidae by the following
derived character states: (1) Adult, male terminalia:
sp2 medially blackish sclerotized (character 53, be-
low); (2) Adult, female terminalia: dorsal margin of
hypogynial valve blackish sclerotized (character 83,
below); (3) Larva: eight anal papillae arranged in four
pairs. The number of anal papillae in other groups of
Tipulidae is either six, four, two, or zero (Gelhaus
1986, Theowald 1957, 1967). As both adult charac-
ters occur also in other genera and subgenera of
Tipulidae, the larval character seems to offer the best
criterion to distinguish Savtshenkia from the remain-
der of Tipulidae.
The systematic position of Savtshenkia within the
Tipulidae is not yet fully established. Savchenko
(1966, 1979, 1983) placed Savtshenkia next to the
subgenus Tipula (Mediotipula Pierre), a relationship
substantiated by the presence in the female pupa of a
laterally placed small tubercle at the apex of the
sheath of the hypogynial valve (see Theowald 1957,
1967). In species of the genus Dolichopeza Curtis, the
sheath of the hypogynial valve is also provided with
an extension, but here the extension is much bigger
than in Mediotipula and Savtshenkia, and is placed
near midlength of the sheath (Byers 1961, Theowald
1957, 1967, Wood 1952). Dolichopeza probably is
not closely related to Mediotipula and Savtshenkia,
and the extension of the sheath of the hypogynial
DID
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
st7 aed spm pmp midv area
o gonst i gonst
aed gd gon fragm
$ ae oe
aed antapod endoph post apod
spmth gen fk st8 st9
hyp viv
Figs. 3-6. Tipula (Savtshenkia) rufina rufina. — 3, male terminalia, lateral view; 4, aedeagal guide with right gonapophysis and
fragmentum, posteromedial view; 5, sperm pump, lateral view; 6, female terminalia, lateral view.
Abbreviations: aed: aedeagus; aed gd: aedeagal guide; ant apod: anterior apodeme; cerc: cercus; comp apod: compressor apo-
deme; endoph: endophallus; fragm: fragmentum; gen fk: genital fork; gon: gonapophysis; goncx: gonocoxite; hyp viv: hypo-
gynial valve; i gonst: inner gonostylus; midv area: midventral area: o gonst: outer gonostylus; post apod: posterior apodeme;
pret: proctiger; sp2: lateral part of genital bridge; spm pmp: sperm pump; spmth: spermatheca; st7 etc.: sternite 7 etc.; sut x:
suture x; tg/ etc.: tergite 7 etc.
valve in Dolichopeza supposedly is not homologous to
that of Mediotipula and Savtshenkia. Theowald
(1978) accepted Savchenko’s views regarding a sister
group relationship of Mediotipula and Savtshenkia.
Another character state which could be indicative of a
close phylogenetic relationship of Mediotipula and
Savtshenkia is the shape of wing cell m1. In both sub-
genera this cell is ‘bell-shaped’, narrowing towards the
wing margin, whereas it usually widens towards the
wing margin in other Tipulidae (character 4, below).
The genital fork in both Mediotipula and Savtshenkia
is broadest at its anterior part, a presumed apomor-
phy within Tipulidae (character 82, below).
276
Mediotipula at present contains 12 recognized species
with a primarily Mediterranean distribution
(Oosterbroek & Theowald 1992).
Conflicting with a presumed sister group relation-
ship of Mediotipula and Savtshenkia is the presence of
a separate sclerite lateral of the gonapophysis in
Savtshenkia. As stated above (section ‘Material, meth-
ods and terminology’), I consider this sclerite homol-
ogous with the so-called fragmentum as found in oth-
er Tipulidae. The presence of a fragmentum is a
derived character state within the Tipulidae and
could substantiate a phylogenetic relationship of
Savtshenkia with the complex of Lunatipula and allies
(here after referred to as the Lunatipula group). The
fragmentum is absent in Mediotipula. The Lunatipula
group contains about 1200 species of Tipulidae, that
is circa one fourth of the total number of species of
the family. Besides the subgenus Lunatipula Edwards,
the Lunatipula group includes the following subgene-
ra of Tipula: Beringotipula Savchenko, Eremotipula
Alexander, Eumicrotipula Alexander, Hesperotipula
Alexander, Labiotipula Alexander, Lindnerina
Mannheims, Odonatisca Savchenko, Pectinotipula
Alexander, Pterelachisus Rondani, Ramatipula
Alexander, Serratipula Alexander, Setitipula
Alexander, Zriplicitipula Alexander, and Vestiplex
Bezzi. The phylogenetic relationships of the species of
the Lunatipula complex are still largely unresolved,
but it is nevertheless clear that not all of the subgene-
ra in their present delimitation represent monophy-
letic taxa (e.g., Lunatipula and Pterelachisus).
In the discussion of the characters, I will refer to
the species of the Lunatipula group and Mediotipula
as outgroups of Savtshenkia.
CHARACTER DISCUSSION
In this section I will discuss the characters used in
the phylogenetic analysis of Savtshenkia. Preceding a
more detailed discussion of each character, a couplet
shortly defines the alternative character states and gi-
ves their codes. The character state matrix is presen-
ted in table 4. Although I did not examine the male of
gimmerthali mattheyi, this subspecies is, according to
its authors, very similar to the nominotypical form,
differing only in minute discriminating characters.
Therefore, I feel safe to copy the score of the male
characters of gimmerthali gimmerthali for this subspe-
cies. In the discussion of the characters below, the re-
sulting strict consensus tree (fig. 2) will often be men-
tioned to point out the species which show a
particular character state. I have done this in order to
save space and for convenience, rather than presen-
ting a list of species for each of the distinguished sta-
tes. The results of the phylogenetic analysis are dis-
cussed in the next section. Species and subspecies of
Savtshenkia which for various reasons are not inclu-
ded in the character discussion below, are dealt with
in Appendix B. A list of autapomorphies of the spe-
cies and subspecies of Savtshenkia is presented in
Appendix C.
Antenna
1. — Flagellum: (0) well developed in male; (1) short,
about the length of head or shorter.
The male antenna of most species of Savtshenkia is
well developed and relatively long, with the flagel-
lomeres being about 2.5 times as long as their diame-
ter at the greatest width, or longer (figs. 7, 11, 171).
H. DE Jone: Phylogeny of Tipula (Savtshenkia)
A similar state represents the presumed groundplan
condition of the Lunatipula group and Mediotipula.
A few members of Savtshenkia are characterized by a
conspicuously short flagellum in the male. The flagel-
lomeres of the pertaining species are about two times
as long as their diameter or shorter (fig. 8). The spe-
cies with exceptionally short male antennae are atlas,
breviantennata, cyrnosardensis, and villeneuvii.
2. — Flagellum: (0) male flagellomeres well
developed; (1) male flagellomeres longer than
standard length.
As noticed under the previous character, the flagel-
lomeres of the males of Savtshenkia are usually about
2.5 times as long as their diameter at greatest width or
slightly longer. Flagellomeres distinctly longer than
this are found in alpha, grisescens, holoptera, pagana,
and most Afrotropical species (the clade elgonensis to
chrysocephala; figs. 9, 10, 152). In the pertaining spe-
cies, the male flagellomeres are about 5 to 6.5 times as
long as wide. The only Afrotropical species of
Savtshenkia of which the male antennae are not con-
spicuously long are hancocki and nephrotomoides (and
the unexamined imperfecta which doubtfully belongs
to Savtshenkia; see Appendix B).
3. — Flagellum: (0) male flagellomeres with slightly
enlarged base, remainder cylindrical; (1)
flagellomeres more or less reniform.
In the genus Tipula sensu lato, the flagellum is usu-
ally composed of eleven segments of which the inter-
mediate ones have a slightly bulbous base and a cylin-
dricalapical parc (ie 101522 F2) iS
character state probably represents the groundplan
condition of the Lunatipula group and Mediotipula.
The same state is also found in most species of
Savtshenkia. The species lundbladi and multipicta dif-
fer from the other species of Savtshenkia by having the
intermediate flagellomeres more or less reniform (fig.
11). In the Lunatipula group, a similar form of the
flagellomeres is present in most species of the subge-
nus Vestiplex. These species appear, however, to rep-
resent a more derived group within Vestiplex.
Wing
4, — Cell m1 shape: (0) widening towards wing
margin; (1) constricted before wing margin.
In the Tipulidae veins M1 and M2, enclosing cell
ml, usually diverge towards the wing margin (fig.
12). This situation probably represents the ground-
plan condition of the Lunatipula group, which con-
tains only a few species that have cell m1 constricted
near the margin of the wing. In most species of
Mediotipula, cell m1 is bell-shaped or paunchy
(‘bauchig’, Mannheims & Pechlaner 1963; Theowald
1973) with veins M1 and M2 approaching towards
Dh
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 7-15. — 7-11, male antenna; 7, Tipula (Savtshenkia) hartigiana; 8, T. (S.) atlas, 9, T. (S.) grisescens, 10, T. (S.) elgonensis,
11, 7. (Sì) lundbladi; 12-15, tip of wing; 12, 7. (S.) grisescens, 13, T. (S.) goriziensis 14, T. (S.) cheethami, 15, T. (S.) akeleyi.
Abbreviations: M1 etc.: medial vein 1 etc.; R4 etc.: radial vein 4 etc.; r-m: radial-medial crossvein.
the margin of the wing, a state which probably repre-
sents the groundplan condition of Mediotipula. The
majority of species of Savtshenkia also show this latter
character state (figs. 13-15). Only a few species of
Savtshenkia have veins M1 and M2 diverging towards
the wing tip, viz. alpha, glaucocinerea, grisescens, and
postposita. The species subnodicornis usually has cell
ml narrowing, but specimens of this species are
known which show a widening cell m1.
278
5. — Cells of wing tip: (0) membrane covered with
microtrichia only; (1) membrane partly covered with
macrotrichia.
Most species of Tipulidae have the membrane of
the wing tip covered with microtrichia only, appear-
ing bare under low magnification (figs. 12, 13, 15).
This situation is found in the species of the
Lunatipula group, Mediotipula, and in most species of
Savtshenkia. The species cheethami and simulans are
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
Figs. 16-31. — 16, Tipula (Savtshenkia) grisescens, aedeagal guide with right gonapophysis and fragmentum, posteromedial
view; 17, T. (S.) hartigiana, aedeagal guide with right gonapophysis and fragmentum, posteromedial view; 18, 7. (S.) atlas,
aedeagal guide with right gonapophysis, posteromedial view; 19, 7. (S.) draconis, aedeagal guide with gonapophyses, ventral
view after microscopic slide holotype; 20, 7. (S.) aberdareica, aedeagal guide with right gonapophysis, posteromedial view; 21,
T. (S.) obsoleta, aedeagal guide with right gonapophysis and fragmentum, posteromedial view; 22, 7: (S.) fragilis, aedeagal gui-
de with right gonapophysis and fragmentum, posteromedial view; 23, 7. (S.) limbata, aedeagal guide with right gonapophy-
sis and fragmentum, posteromedial view; 24, 7. (S.) pechlaneri, aedeagal guide with right gonapophysis and fragmentum, pos-
teromedial view; 25-27, 7. (S.) haennii, 25, aedeagal guide, tip, posterior view; 26, right gonapophysis, medial view; 27, right
fragmentum, posteromedial view; 28-30, T. (S.) corsosignata, 28, aedeagal guide, tip, posterior view; 29, right gonapophysis,
medial view; 30, right fragmentum, posteromedial view; 31, 7. (S.) subsignata subsignata, right gonapophysis, medial view.
unique within Savtshenkia by having macrotrichia on
the membrane of the wing tip. The macrotrichia cov-
er the wing to a larger extent in simulans than in chee-
thami. The density of the macrotrichia is also higher
in the former species. Macrotrichia are usually re-
stricted to cells r4, r5, and ml in cheethami (fig. 14),
while in sémulans cell m2 and the tip of cell m3 are al-
so provided with macrotrichia.
6. — Crossvein r-m: (0) apical section directed
towards wingtip, or at most transverse to
longitudinal axis of wing; (1) apical section directed
towards base of wing.
In Tipulidae crossvein r-m usually lies in an
oblique position relative to the longitudinal axis of
the wing, with its apical part directed towards the
wing tip. This situation is found in the species of the
Lunatipula group, Mediotipula, and in the majority of
species of Savtshenkia (figs. 12-14). Three species of
Savtshenkia have the apical end of r-m directed to-
wards the base of the wing, viz. akeleyi, caligo, and
chrysocephala (fig. 15).
29,
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Male terminalia
7.— Aedeagal guide, apex: (0) tapering towards tip;
(1) at tip with flaring rim.
As is usual in the Tipulidae, the apex of the aedea-
gal guide is the narrowest part of this structure in the
species of the Lunatipula group, Mediotipula, and the
majority of species of Savtshenkia (figs. 4, 16, 17, 19,
20, 22-25, 28, 162, 182, 194). The species atlas,
breviantennata, and obsoleta have the apex of the ae-
deagal guide bent outwards as a flaring rim (figs. 18,
ZIE
8. — Aedeagal guide, apex: (0) gradually tapering
towards tip; (1) subapically abruptly narrowed,
terminating in a slender point.
The aedeagal guide in Tipulidae normally gradual-
ly narrows towards its apex, as it does in almost all
species of Savtshenkia (caudal view, fig. 25). The apex
of the aedeagal guide of corsosignata, subsignata sub-
signata, and subsignata cazorla differs from that of the
other species of Savtshenkia by the abruptly constrict-
ed apex which terminates in a slender tip (fig. 28).
9. — Aedeagal guide, gonapophysis: (0) well
developed; (1) very short or almost completely
absent.
Gonapophyses are usually well developed in the
Tipulidae, a state which probably represents the
groundplan condition of the Lunatipula group and
Mediotipula. The majority of species of Savtshenkia
also have well developed gonapophyses which occur
in a wide range of sizes and shapes (figs. 4, 17-24, 26,
29, 31, 182, 194). The gonapophyses are very short
or almost completely absent in the species alpha, alpi-
UM, boreosignata, grisescens, interserta, and macarones-
ica (figs. 16, arrow, 162).
10. — Aedeagal guide, gonapophysis: (0) dorsal
margin evenly fused with base of aedeagal guide; (1)
anterodorsally slightly extended, thus with free-lying
dorsal tip.
In general, the dorsal margin of the gonapophysis
in the Tipulidae gradually merges with the base of the
aedeagal guide. This situation probably represents the
groundplan condition of the Lunatipula group and
Mediotipula. The gonapophyses of most species of
Savtshenkia also show this character state (figs. 16,
21-24, 162, 182, 194). The species of the clade har
tigiana to simulans are characterized by their typically
shaped gonapophyses, which differ from the gona-
pophyses of the other species of Savtshenkia by the
presence of a free anterodorsal extension (figs. 17, ar-
row, 18).
11. — Aedeagal guide, gonapophysis: (0) pubescent
or short haired; (1) long haired.
280
The covering of the gonapophyses in the Tipulidae
usually consists of pubescence or short hairs, as is the
presumed groundplan state of the Lunatipula group
and Mediotipula. A similar condition is found in the
majority of species of Savtshenkia (figs. 4, 17-19, 21-
24). A number of Afrotropical species of Savtshenkia
are distinguished from the remainder of the subgenus
by the dense and elongate hairing of the gonapophy-
ses, viz. aberdareica, akeleyt, asbolodes, caligo, chrysoce-
phala, and hancocki (fig. 20).
12. — Aedeagal guide, gonapophysis: (0) present,
variously shaped; (1) a large and elongate structure.
The gonapophyses of the Tipulidae show a wide
range of shapes. When present, they are usually well
developed and rather slender. This is the state that oc-
curs in most of the species of Savtshenkia and is the
presumed groundplan condition of its outgroups (-
figs. 4, 17-21, 23, 24, 182, 194). The Nearctic species
fragilis and phoroctenia differ from the majority of
species of Savtshenkia by their extremely large and el-
ongate gonapophyses (fig. 22). The only other species
of Savtshenkia with similarly enlarged gonapophyses
is the western Mediterranean cyrnosardensis.
13. — Aedeagal guide, gonapophysis: (0) a relatively
slender structure; (1) a large blade.
As noted under the previous character, the plesio-
morphous condition of the gonapophyses in
Savtshenkia is probably represented by a well devel-
oped, rather slender structure (figs. 4, 17-21, 23, 24,
182, 194). Large mediolaterally flattened gonapophy-
ses are found in corsosignata, haennii, subsignata sub-
signata, and subsignata cazorla (figs. 26, 29, 31; see al-
so next character).
14. — Aedeagal guide, gonapophysis: (0) at apex at
most moderately enlarged; (1) dorsally and ventrally
extended and thus widening towards broad apex.
The two subspecies subsignata subsignata and sub-
signata cazorla are characterized by their expanded
gonapophyses (fig. 31). This character state corre-
sponds with character 6 of Dufour (1991).
15. — Aedeagus: (0) short, directly curving from
sperm pump to aedeagal guide, anteriorly reaching
as far as abdominal segment 8; (1) elongate,
anteriorly running to segment 7 or beyond.
A relatively short aedeagus which loops in the me-
dian sagittal plane from the sperm pump through ab-
dominal segment 8 and from there to the aedeagal
guide, is the presumed groundplan state of this fea-
ture in the Lunatipula group. The species of
Mediotipula also show this character state, as do a
number of species of Savtshenkia (figs. 3, 155). Other
species of Savtshenkia have a longer aedeagus. The ae-
deagus in these species reaches anteriorly at least to
segment 7 and often lies coiled inside the abdomen (-
figs. 173, 185). The species involved constitute the
clades confusa to aster and fragilis to omega. Extremely
long aedeagi were found in corsosignata, ornata, and
staegeri, species in which the aedeagus reaches abdom-
inal segment 1. The species pair boreosignata and ele-
onorae is characterized by a peculiarly modified aedea-
gal complex (character 52). Although belonging to
the clade fragilis to omega, they have a short aedeagus.
16. — Fragmentum: (0) either absent, or posterior
margin convex or slightly concave; (1) posterior
margin subapically with sharp emargination.
The fragmenta of the species of the Lunatipula
group and Savtshenkia occur in a diversity of forms.
Usually, the posterior margin near the dorsal tip of
the structure is convex. This is the presumed ground-
plan condition of the Lunatipula group and the com-
mon character state in Savtshenkia (figs. 4, 16, 17, 22-
24, 27, 30, 162, 182, 194). (The fragmentum is
absent in the species of Mediotipula.) The fragmenta
of the species holoptera, obsoleta, and pagana differ
from the standard situation of Savtshenkia by the
presence of a distinct emargination just below the
dorsal apex (fig. 21, arrow).
17. — Fragmentum: (0) either absent, or anterior
part sclerotized as the remainder of the structure; (1)
anterior part dorsally blackish sclerotized.
In general, the fragmentum is yellowish-brown in
colour and uniformly sclerotized in the species of the
Lunatipula group and Savtshenkia (figs. 4, 16, 17, 21,
22, 24, 162, 182, 194). A unique state is found in the
species fragilina, limbata, and subvafra, where the
dorsal margin of the anterior part of the fragmentum
is blackish sclerotized (fig. 23).
18. — Fragmentum: (0) either absent, or anterior
part a rounded lobe; (1) anterior part a separate
anterocaudally flattened extension with slender tip.
The fragmentum consists of a single undivided
sclerite in the species of the Lunatipula group and
most species of Savtshenkia (figs. 4, 16, 17, 21-24,
162, 182, 194). In corsosignata, haennii, subsignata
subsignata, and subsignata cazorla, the anterior part of
the fragmentum is characteristically modified in a
slender tipped structure which stands apart from the
posterior part (figs. 27, 30). The few hairs of the
brush on top of the anterior part of the fragmentum
are very short in these four taxa.
19. — Gonocoxite, midventral area: (0) variously
shaped, usually tumescent; (1) with cone-shaped
membranous extension.
The midventral area in between the gonocoxites is
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
represented by a tumescent structure in the presumed
groundplan state of the Lunatipula group and
Mediotipula. The majority of species of Savtshenkia
also have a tumescent midventral area (fig. 32), but a
number of species show a modification of this part of
the terminalia. The species atlas, breviantennata, chee-
thami, hartigiana and simulans have the midventral
area provided with a ventrally directed cone-shaped
membranous projection. The projection is relatively
low in atlas and breviantennata (fig. 34), in the other
species listed it is longer and more slender (figs. 33,
35). The structure is placed near the posteroventral
margin of the gonocoxites in cheethami (fig. 35), in
the four other species it has a more anterior position
(figs. 33, 34).
Other species of Savtshenkia with a projection to
the area in between the gonocoxites are alpium, aspro-
montensis, macaronesica, ornata, phoroctenia, and a
few Afrotropical species. As the actual shapes of the
projections differ notably among these species, a
number of different character states are distinguished.
In the species alpium and macaronesica, the append-
age is membranous, dorsoventrally flattened and
midventrally placed (character 21; fig. 39), in ornata
it is swollen, sclerotized and posteriorly projecting
(fig. 40), in phoroctenia it is slender, sclerotized and
posteriorly projecting (fig. 41). The modifications of
the midventral area in ornata and phoroctenia are
characteristic of the respective species and represent
autapomorphies (see Appendix C). Males of the spe-
cies aspromontensis originating from Greece
(Peloponnisos, Crete) have the midventral area poste-
riorly terminating in a small and low extension. A
similar extension is absent in specimens of aspro-
montensis from the type locality in southern Italy. I
could not find other distinguishing characters
between male specimens of the populations from
Greece and from Italy and consider them conspecific.
The Afrotropical species of Savtshenkia with a modi-
fied midventral area include akeleyi, caligo, chrysoce-
phala, and nephrotomoides. These species are charac-
terized by a relatively large and bulbous extension to
the midventral area (next character).
20. — Gonocoxite, midventral area: (0) posterior part
not bulbous; (1) bulbous.
As noted under the previous character, the midven-
tral area in Savtshenkia usually is tumescent. The
Afrotropical species caligo, chrysocephala, nephrot-
omoides, and, to a lesser extent, akeleyi, are distin-
guished by a posteroventrally projecting lobe near the
posterior margin of the midventral plate. The lobe is
large in the first three species listed (fig. 36), in akeleyi
it is relatively low (fig. 38).
21. — Gonocoxite, midventral area: (0) without
281
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 32-45. — 32-41, male sternite 9, left gonocoxite and midventral area, lateral view; 32, Tipula (Savtshenkia) grisescens 33,
T. (Sì) hartigiana; 34, T. (S.) atlas 35, T. (S.) cheethami; 36, T. (S.) nephrotomoides, 37, T. (S.) aberdareica; 38, T. (S.) ake-
ley, 39, T. (S.) alpium; 40, T. (S.) ornata; 41, T. (S.) phoroctenia, 42-45, male sternite 9, gonocoxites and midventral area,
ventral view; 42, 7. (S.) baltistanica; 43, T. (S.) lundbladi; 44, T. (S.) nielsent 45, T. (Sì) haennii.
Abbreviations: midv area: midventral area; sut x: suture x.
flattened appendage; (1) at about midlength with
dorsoventrally flattened appendage.
As stated above, the midventral area in between the
gonocoxites is tumescent in most species of
Savtshenkia. The species alpium and macaronesica are
distinguished from the other species of Savtshenkia by
the presence of a dorsoventrally flattened membra-
282
nous appendage at about midlength of the midventral
area. The appendage is long in alpium (fig. 39), in
macaronesica it is considerably shorter. (The unexam-
ined subalpium has a small tongue-shaped appendage
to the midventral area and is probably closely related
to alpium and macaronesica; see Appendix B).
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
Figs. 46-51. Male tergite 9-10 and dorsal part of left gonocoxite plus appendages, lateral view. — 46, Tipula (Savtshenkia) har-
tigiana; 47, T. (S.) tulipa; 48, T. (S.) haennii, 49, T. (S.) boreosignata; 50, T. (S.) benesignata; 51, T. (S.) persignata tofina.
22. — Gonocoxite, midventral area: (0) broad and
usually separated from gonocoxites by membranous
zone; (1) narrow and fused with gonocoxites.
The midventral area is probably broad in the
groundplan states of the Zunatipula group and
Mediotipula. The majority of species of Savtshenkia
also have a broad midventral area in between the gon-
ocoxites (figs. 42-44). It either is membranous, or
partly or entirely sclerotized. In most Savtshenkia spe-
cies with a sclerotized midventral area, a membranous
zone (partly) separates the plate from the gonocoxites.
The species pechlaneri and the members of the clade
haennii to omega have a slender parallel sided and
sclerotized midventral area which is largely fused with
the ventromedial margins of the gonocoxites (fig. 45).
23. — Gonocoxite, posterodorsal margin: (0) with
scattered short setae; (1) with concentration of long
and strong setae.
The gonocoxites of most species of Tipulidae are
covered with scattered setae, a situation which prob-
ably represents the groundplan condition of this char-
283
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
us lat rdg post pt
Figs. 52-63. Left inner gonostylus. — 52, Tipula (Savtshenkia) grisescens, dorsal view; 53, 7. (S.) villeneuvii, dorsal view; 54, 7:
(S.) goriziensis, dorsal view; 55, 7. (S.) rufina rufina, lateral view; 56, 7: (S.) aberdareica, lateral view; 57, T. (S.) akeleyi, lateral
view; 58, 7. (S.) confusa, lateral view; 59, 7. (S.) ornata, lateral view; 60, 7. (S.) serrulifera, lateral view; 61, 7. (S.) aspromon-
tensis, lateral view; 62, 7. (S.) fragilis, medial view; 63, 7. (S.) odontostyla, medial view.
Abbreviations: ant pt: anterior part; lat rdg: lateral ridge; post pt: posterior part.
acter for the Lunatipula group and Mediotipula. It is
also found in most species of Savtshenkia (figs. 3, 46,
48, 49, 155). Within Savtshenkia, the species of the
clades tulipa to simulans and benesignata to omega are
distinguished by a concentration of strong setae along
the posterodorsal margin of the gonocoxites (figs. 47,
arrow, 50, 51, 173, 185). The setae are placed in two
or three irregular rows. They extend dorsally towards
the membranous suture separating the gonocoxite
from tergite 9-10 in the species of the clade tulipa to
simulans and in benesignata (figs. 47, 50). In the spe-
cies of the clade persignata to omega, the setae are con-
fined to a more ventral position (fig. 51, 173, 185).
24. — Gonocoxite, posterodorsal margin: (0)
posterodorsal corner rounded off; (1) with posterior
projection.
The posterodorsal corner of the gonocoxites is pre-
sumably rounded off in the groundplan condition of
the Lunatipula group, as it is in the majority of species
of Savtshenkia. In the species of Mediotipula and a
number of species of Savtshenkia, the posterodorsal
margin of the gonocoxite is extended into a more or
less well developed projection (fig. 48, arrow). The
species of Savtshenkia which show this character state
284
are those of the range fragilina to subsignata cazorla.
25. — Gonocoxite, suture x: (0) angle between suture
x and the median line about 45° or less; (1) angle
between suture and median line much wider, up to
90°.
Suture x usually stands oblique to the median line
of the body under an angle of about 45° or less in the
Tipulidae. This probably is the groundplan condition
of the character for the Lunatipula group and
Mediotipula, and is the general state in Savtshenkia (-
figs. 42, 44, 45). In the species /undbladi, multipicta,
rufina, and the Afrotropical members of the subge-
nus, the angle is much wider, often reaching a magni-
tude of about 90° (fig. 43). (The direction of suture x
could not be adequately studied in the celluloid prep-
aration of the holotype of e/gonensis, the only known
specimen of this species.)
26. — Gonocoxite, suture x: (0) narrow; (1) broad.
Suture x usually is rather narrow in the Tipulidae,
a condition considered the groundplan state of the
Lunatipula group and Mediotipula. A narrow suture x
is also found in the majority of species of Savtshenkia
(figs. 3, 32-39, 41-44). The species and subspecies
corsosignata, haennii, subsignata subsignata, and sub-
signata cazorla differ from the other members of
Savtshenkia by their broad suture x (fig. 45, arrow).
The suture is straight in these four taxa. Other species
of Savtshenkia with a broad suture x are fragilis and
ornata. \n fragilis, suture x is sigma-shaped, in ornata
it is, as many other genital features in this species,
very different from that of the remainder of species of
Savtshenkia. In ornata, suture x encompasses an iso-
lated sclerite (fig. 40, arrow; see Appendix C).
. 27. — Inner gonostylus, anterior part: (0) ventral
surface smooth, or at most rugged; (1) serrate.
The ventral surface of the anterior part of the inner
gonostylus usually is smooth in Tipulidae. It is con-
sidered the groundplan state of the Zunatipula group
and Mediotipula, and is found in the majority of spe-
cies of Savtshenkia (all figures that depict the inner
gonostylus in side view, except 57). Three
Afrotropical members of Savtshenkia are character-
ized by a partly serrate lower margin of the anterior
part of the inner gonostylus, viz. akeleyi, caligo, and
chrysocephala (fig. 57, arrow).
28. — Inner gonostylus, anterior part: (0) crest absent
on posterodorsal margin of anterior part; (1) crest
present.
The anterior part of the inner gonostylus probably
lacks a crest in the groundplan state of the Lunatipula
group. All species of Mediotipula are devoid of a crest
on this part, as are most species of Savtshenkia (the
majority of figures depicting the inner gonostylus).
Three species of Savtshenkia have a well developed
membranous crest on the posterodorsal margin of the
anterior part of the inner gonostylus, viz. aspromonte-
nsis, aster, and serrulifera. The crest is dorsally round-
ed in aster and serrulifera (fig. 60), but it is grotesque-
ly widening towards its slightly concave dorsal margin
in aspromontensis (fig. 61).
29. — Inner gonostylus, anterior part: (0) anteriorly
terminating in various ways; (1) anteriorly
terminating in a prolonged nipple.
The anterior part of the inner gonostylus shows a
wide range of forms in the Lunatipula group,
Mediotipula, and Savtshenkia. In Savtshenkia it is of-
ten short and terminates anteroventrally in a more or
less acute tip (the majority of figures showing the in-
ner gonostylus). The species aspromontensis, aster, and
serrulifera have a uniquely shaped anterior part which
gradually tapers towards an anteriorly extended slen-
der nipple-like projection (figs. 60, 61). The exten-
sion is longest in aspromontensis (fig. 61) and aster.
30. — Inner gonostylus, anterior part: (0) tip single;
(1) tip bifid.
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
The anterior part of the inner gonostylus termi-
nates in a single tip in the majority of Tipulidae, a sit-
uation representing the presumed groundplan condi-
tion of the Lunatipula group and Mediotipula. The
majority of species of Savtshenkia also have the tip of
the anterior part of the inner gonostylus undivided (-
figs. 55-63, 68, 70-75, 161, 181, 193). The species
corsosignata, nielseni, and the representatives of the
clade fragilina to gimmerthali differ from the other
species of Savtshenkia in having a bifid anterior tip of
the inner gonostylus. The tip in most of these species
shows a relatively broad dorsal and finer lateroventral
point (figs. 64-67). In corsosignata (fig. 69), the small-
er tip lies in a more lateral position compared with
that of the other species with a bifid tip.
31. Inner gonostylus, anterior part: (0) narrowing
towards pointed tip; (1) abruptly bent, almost
truncate at tip.
The majority of species of Savtshenkia are charac-
terized by a gradually curved and pointed tip of the
anterior part of the inner gonostylus (lateral view; see
figs. 55-72). The same situation probably is the
groundplan condition of the Lunatipula group and
Mediotipula. Within Savtshenkia, the species of the
clade persignata to omega are distinguished from the
remainder of the subgenus by the broad, almost par-
allel-sided anterior part of the inner gonostylus which
terminates in a rather steep front end (figs. 73-75,
1315, 193):
32. — Inner gonostylus, base: (0) posteriorly
rounded; (1) posteromedially terminating in pointed
extension.
Usually, the base of the inner gonostylus is round-
ed at its posterior margin in the Tipulidae, a situation
representing the presumed groundplan condition of
the Lunatipula group and Mediotipula. It is also
found in most species of Savtshenkia (dorsal view, fig.
52, arrow). A number of species of Savtshenkia are
distinguished by the presence of an acute point at the
posteromedial corner of the base of the inner gonos-
tylus (figs. 53, arrow, 54). The species concerned con-
stitute the clade hartigiana to simulans.
33. — Inner gonostylus, lateral ridge, ventral part: (0)
without extensions; (1) with one or two acute
blackish pointed extensions.
The ventral part of the lateral ridge of the inner go-
nostylus gradually merges with the main body of the
inner gonostylus in most species of Savtshenkia. The
three species cheethami, goriziensis, and simulans differ
from the remainder of species of Savtshenkia by the
presence of one or two acute blackish points at the
end of the lower part of the lateral ridge of the inner
gonostylus. The points are located near midheight of
285
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 64-75. Left inner gonostylus, lateral view. — 64, Tipula (Savtshenkia) limbata; 65, T. (S.) pechlaneri, 66, T. (S.) gim-
merthali gimmerthali, 67, T. (S.) gimmerthali pteromaculata; 68, T. (S.) haennii, 69, T. (S.) corsosignata; 70, T. (S.) subsignata
subsignata; 71, T. (S.) boreosignata; 72, T. (S.) benesignata; 73, T. (S.) persignata tofina; 74, T. (S.) signata; 75, T. (S.) man-
nheimsi.
the body of the inner gonostylus. In goriziensis, a sin-
gle point is present (fig. 54, arrow), while cheethami
and simulans have two points, the posterior of which
is bifid in simulans.
34. — Inner gonostylus, lateral ridge: (0) ventrally
directed and terminating at about midlength of base
of inner gonostylus; (1) in ventral part angularly
286
bent and anteriorly extended, terminating in front of
base of inner gonostylus.
The lateral ridge of the inner gonostylus is ventral-
ly directed in the majority of species of Savtshenkia.
The lateral ridge of subsignata subsignata and subsig-
nata cazorla is present as a sharp edge directed to-
wards the anterior margin of the inner gonostylus.
The ridge makes a distinct angle in its posteroventral
section near the base of the inner gonostylus (fig. 70).
Dufour (1991, character 3) recognized the pres-
ence of a ridge, on what he referred to as the median
part of the inner gonostylus, as a synapomorphy of
the subspecies subsignata subsignata and the then
newly described subsignata cazorla and subsignata
haennii. Furthermore, Dufour (1991, character 4)
considered the continuity of the ventral and posterior
parts of the ridge, as found in subsignata subsignata
and subsignata cazorla, a synapomorphy of these two
subspecies. Regarding the shape of the ridge in other
species of Savtshenkia, it appears, however, that a con-
tinuous ridge should be judged a plesiomorphy in
subsignata subsignata and subsignata cazorla.
Consequently and contrary to Dufour, I regard the
separation of the posterior and ventral parts of the
ridge in haennii an autapomorphy of that species (fig.
68; see Appendix C).
35. — Inner gonostylus, posterior part: (0) variously
shaped; (1) posterolaterally extended, cup-shaped,
laterodorsal margin blackened and serrate.
The groundplan condition of the posterior part of
the inner gonostylus in the Lunatipula group and
Mediotipula is probably represented by a relatively
simple and single structure. The posterior part is
rounded and little developed in most species of
Savtshenkia. The species confusa, eugeni, and jeekeli
differ from the other species of Savtshenkia by the
configuration of the posterior part. In these three spe-
cies, it projects posterolaterally and carries acute pro-
cesses along its blackish sclerotized laterodorsal mar-
gin (fig. 58). The species confusa has a serrate and
blackened rim running medially of and parallel to the
posterolateral margin (fig. 58; see Appendix C).
36. — Inner gonostylus, posterior part: (0) variously
shaped; (1) a posteriorly projecting, often rather
acute, structure.
As stated above, the posterior part of the inner go-
nostylus is moderately developed and rounded off in
most species of Savtshenkia. The members of the
clade odontostyla to omega are characterized by a more
caudal development of the posterior part. In these
species, the posterior part either is a rather slender
and often somewhat acute structure (fig. 63), or a
more enlarged and posteriorly produced extension
(figs. 68-75, 181, 193; see also next character). A cau-
dally produced posterior part of the inner gonostylus
is also present in the species ornata (fig. 59).
37. — Inner gonostylus, posterior part: (0) a relatively
small projection; (1) posteriorly extended, large.
As mentioned under the previous character, the
posterior part of the inner gonostylus is usually little
developed in the species of Savtshenkia. The species
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
benesignata, mannheimsi, omega, persignata, signata,
and trinacria are distinguished among the species of
the subgenus by a strikingly large posterior part which
is of about the same length as the anterior part (figs.
72-75, 181, 193). The posterior part in these species
is vertically orientated and lies more or less in line
with the anterior part.
38. — Inner gonostylus, posterior part: (0) dorsal
surface even; (1) dorsal surface corrugated.
The dorsal surface of the posterior part of the inner
gonostylus is even in the majority of species of
Savtshenkia, as is the probable groundplan condition
of the Lunatipula group and Mediotipula. Within
Savtshenkia the dorsal surface of this part of the inner
gonostylus is corrugated in both persignata persignata
and persignata tofina (fig. 73).
39. — Inner gonostylus, posterior part: (0) variously
shaped; (1) anterior section anterodorsally extended
towards anterior part of inner gonostylus.
As noted under character 37, a number of species
of Savtshenkia have a caudally produced and large
posterior part of the inner gonostylus. The extended
posterior part of the inner gonostylus is low in its an-
terior half in benesignata, persignata persignata, and
persignata tofina (figs. 72, 73), as it is in the other spe-
cies of Savtshenkia. In the species mannheimsi, omega,
signata, and trinacria it is anterodorsally protruding
(figs. 74, 75, 181, 193).
40. — Inner gonostylus: (0) variously shaped; (1)
similarly shaped, with high anterior part and
blackish sclerotized triangular projection
posterolaterally.
The inner gonostylus shows a wide variety of forms
throughout Savtshenkia. The two subspecies rufina
rufina and rufina maderensis display a remarkable
similarity in the structure of the inner gonostylus
which differs most notably from that of other species
of Savtshenkia by the shape of the anterior part and
the location of the posterolateral projection (fig. 55).
41. Inner gonostylus: (0) variously shaped; (1)
similarly shaped, anterior part anterodorsally
straightened, gradually narrowing towards bifid tip.
Throughout Savtshenkia a wide variety of forms of
the inner gonostylus can be found. The three recog-
nized subspecies of gimmerthali can be separated from
the other species of the subgenus by the unique shape
the inner gonostylus, of which the anterior part grad-
ually narrows towards a bifid tip (figs. 66, 67).
42. — Inner gonostylus: (0) variously shaped; (1)
similarly shaped, with long sloping caudodorsal
margin.
287
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 76-99. — 76-88, left outer gonostylus, lateral view; 76, Tipula (Savtshenkia) grisescens 77, T. (S.) hartigiana; 78, T. (S.)
lundbladi; 79, T. (S.) multipicta; 80, T. (S.) jeekeli; 81, T. (S.) aspromontensis 82, T. (S.) fragilis, 83, T. (S.) gimmerthali gim-
merthali, 84, T. (S.) gimmerthali pteromaculata; 85, T. (S.) nielseni; 86, T. (S.) sardosignata; 87, T. (S.) signata; 88, T. (S.) man-
nheimst, 89-99, sperm pump and appendages; 89, 7. (S.) subnodicornis, lateral view; 90, 7. (S.) baltistanica, lateral view; 91-
93, 7. (S.) alpium, 91, lateral view; 92, compressor apodeme, posterior view; 93, anterior view; 94-96, 7. (S.) jeekeli, 94,
lateral view; 95, compressor apodeme, posterior view; 96, anterior view; 97, 7. (S.) confusa, lateral view; 98, 7. (S.) graciae, la-
teral view; 99, 7. (S.) boreosigata, lateral view.
The species boreosignata and eleonorae are distin-
guished among the species of Savtshenkia by their
uniquely shaped and very similar inner gonostyli. The
overall structure is rather long, with a well developed
anterior part and a long caudodorsal margin which
gradually slopes towards the low posterior part (fig.
71).
43. — Outer gonostylus: (0) anterior margin convex
288
or slightly and evenly concave; (1) concave, with
distinct bend at about two-fifth of length from base.
The anterior margin of the outer gonostylus is ei-
ther entirely convex or evenly concave in its lower half
in the presumed groundplan condition of the
Lunatipula group and Mediotipula, and in the major-
ity of species of Savtshenkia (figs. 76, 78-88, 160,
180, 192). The three species atlas, breviantennata,
and hartigiana have the anterior margin of the outer
gonostylus angularly concave near two-fifth of its
height (fig. 77, arrow).
44. — Outer gonostylus: (0) upright structure; (1)
inclined forward.
The outer gonostylus in Tipulidae usually is an up-
right structure, as is the presumed groundplan condi-
tion of the Lunatipula group and Mediotipula. The
majority of species of Savtshenkia are also distin-
guished by the presence of a vertically orientated out-
er gonostylus (figs. 76, 77, 80, 82-88, 160, 180, 192).
The outer gonostylus of the species aspromontensis,
lundbladi, and multipicta leans over anteriorly (figs.
78, 79, 81). The actual shapes of the outer gonostyli
of the three pertaining species differ notably. The
base of the outer gonostylus is low and elongate in
lundbladi (fig. 78) and multipicta (fig. 79), whereas it
is rather short in aspromontensis (fig. 81).
45. — Outer gonostylus: (0) anterior part rather
broad; (1) slender and elongate.
In general, the outer gonostylus is present as a sin-
gle oblong and rather broad structure in the
Tipulidae, a situation which probably represents the
groundplan condition of this character for the
Lunatipula group and Mediotipula. A broad outer go-
nostylus is found in about half of the species of
Savtshenkia (figs. 76-81, 160). The species of the
clade fragilis to omega are distinguished by the pres-
ence of a rather slender and elongate anterior sub-
structure of the outer gonostylus (figs. 82, 83, 85-88,
180, 192). A similar outer gonostylus with slender
anterior section is found furthermore in the species
eugeni and ornata. The subspecies gimmerthali pte-
romaculata, which is included in the clade fragilis to
omega, has a relatively wide anterior part of the outer
gonostylus (fig. 84).
46. — Outer gonostylus: (0) base variously shaped,
usually relatively narrow; (1) broad, laterally concave
and blackish sclerotized.
As noticed under the previous character, the outer
gonostylus generally is an oblong structure in the
Tipulidae. The base of the outer gonostylus is usually
relatively narrow compared with the length of the
structure, a situation present in the majority of spe-
cies of Savtshenkia (figs. 76, 77, 80, 82-84, 87, 88,
160, 180, 192). The species nielseni (fig. 85) and sar-
dosignata (fig. 86) differ from the remainder of
Savtshenkia by the presence of a large base of the out-
er gonostylus. The base is laterally concave, partly
blackish sclerotized and carries a strong posterior pro-
jection.
47. — Sperm pump, endophallic lumen: (0) bulbous,
ventrally expanded; (1) small.
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
Throughout the Tipulidae, the sperm pump has a
rather large and bulbous endophallic lumen, a situa-
tion which probably represents the groundplan con-
dition of the Lunatipula group and Mediotipula.
Within Savtshenkia a similar condition is found in the
species alpha, grisescens, ignobilis, subnodicornis, and
the species of the clade hartigiana to simulans (figs.
89, 163). The majority of species of Savtshenkia,
however, have an endophallic lumen that is hardly
ventrally expanded (figs. 5, 90, 91, 94, 97, 98, 183,
195). The species concerned include the members of
the clade baltistanica to omega.
48. — Sperm pump, compressor apodeme: (0) a
flattened blade; (1) posteriorly provided with U-
shaped or V-shaped brace.
The groundplan condition of the compressor apo-
deme of the Lunatipula group and Mediotipula prob-
ably consists of a flattened blade with a more or less
distinct dorsomedial emargination. Within
Savtshenkia a similar compressor apodeme is found in
the species that range from alpha to macaronesica (-
figs. 92, 93, 164). The other species of Savtshenkia are
distinguished by the presence of a U-shaped or V-
shaped support on the posterior side of the compres-
sor apodeme (fig. 95, arrow, 96, 184, 196). The spe-
cies involved constitute the clade jeekeli to omega.
49. — Sperm pump: (0) ventral wall convex; (1)
ventral wall flattened or concave.
In Tipulidae, the body of the sperm pump usually
is a bulbous structure which has a convex ventral wall.
This situation probably reflects the groundplan con-
dition of the Lunatipula group and Mediotipula and is
found in about two-fifth of the species of Savtshenkia
(the range alpha to macaronesica; frontal view, figs.
93, 164). Within Savtshenkia the bulb of the sperm
pump has a flat or concave ventral wall in most spe-
cies of the clade jeekeli to omega (figs. 96, 184, 196).
There are, however, a few species within this latter
clade which have a convex ventral wall of the sperm
pump, viz. aspromontensis, eugent, and interserta.
50. — Sperm pump, posterior apodemes: (0) well
developed and posteriorly or posterodorsally
directed; (1) elongate and anteriorly extended.
In Tipulidae, the posterior apodemes of the sperm
pump are generally well developed, rather broad and
posteriorly or posterodorsally directed. This character
state probably represents the groundplan condition of
the Lunatipula group and Mediotipula. The majority
of species of Savtshenkia also display this character
state (figs. 5, 89-91, 94, 163, 183, 195). In contrast,
the members of the clade confusa to aster are distin-
guished by the presence of elongate and anteriorly ex-
tended posterior apodemes. The tips of the posterior
289
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 100-112. — 100, 101, sp2 and part of tergite 9-10, posteromedial view; 100, Tipula (Savtshenkia) odontostyla; 101, T.
(S.) limbata; 102-112, male sternite 8, posterior margin; 102, 7: (S.) grisescens, ventral view; 103, 7. (S.) lundbladi, ventral
view; 104, 7. (S.) rufina rufina, posterior view; 105, 7. (S.) alpium, ventral view; 106, 7. (S.) obsoleta, ventral view; 107, 7.
(S.) holoptera, ventral view; 108, T. (S.) confusa, ventral view; 109, T. (S.) aster, ventral view; 110, 7: (S.) limbata, ventral view;
111, 7. (S.) pechlaneri, ventral view; 112, 7. (S.) invenusta invenusta, ventral view.
Abbreviations: sp2: lateral part of genital bridge; tg9-10: tergite 9-10.
apodemes in these species lie alongside the compres-
sor apodeme (figs. 97, 98).
51. — Sperm pump, posterior apodemes: (0) well
developed, broad; (1) slender.
As noticed under the previous character, the poste-
rior apodemes of the majority of species of
Savtshenkia are well developed and broad (figs. 5, 89-
290
91, 94, 98, 163, 183, 195). Three species are distin-
guished from the remainder of the subgenus by their
long and slender posterior apodemes, viz. confusa,
interserta, and ornata (fig. 97). The posterior apo-
demes of these three species are anteriorly extended
(see previous character).
52. — Sperm pump: (0) general shape of the standard
tipulid type; (1) general shape different, partly
heavily sclerotized.
The sperm pump in Tipulidae usually consists of a
bulbous main body, which is surrounded in the hori-
zontal plane by a pair of anterior and a pair of poste-
rior apodemes, and perpendicular and dorsal to these
a single compressor apodeme. The anterior and pos-
terior apodemes are attached to the main body of the
sperm pump at about midheight and the aedeagus
leaves the sperm pump in between the anterior apo-
demes. This configuration doubtlessly represents the
groundplan condition of the Lunatipula group and
Mediotipula. The majority of species of Savtshenkia
also show a sperm pump of this form (figs. 5, 89-98,
163, 164, 183, 184, 195, 196). However, the species
boreosignata and eleonorae possess a peculiarly built
sperm pump, which has the heavily sclerotized main
body lying just dorsal of (boreosignata, fig. 99) or on
(eleonorae) a membranous disc-like structure which
contains the remnants of the anterior and posterior
apodemes. The heavily sclerotized compressor apo-
deme seems to be fused with the main body of the
pump. In boreosignata, the short aedeagus appears to
be attached to the ventral side of the main body, after
which it makes a loop at the level of the disc-like
structure containing the anterior and posterior apo-
demes. Compared with the sperm pump of the other
species of Savtshenkia, the whole structure is small in
both boreosignata and eleonorae.
53. — Sp2: (0) uniformly and moderately sclerotized;
(1) medially blackish sclerotized.
Throughout the Tipulidae, sp2 generally is uni-
formly sclerotized. This situation is found in the
Lunatipula group and in Mediotipula and represents
the groundplan state of this character for both taxa.
All species of Savtshenkia as recognized in this paper
have the medial margin of sp2 blackish sclerotized
(figs. 100, 101, 159, 179, 191). The only other
Tipulidae in which a similar situation was studied are
species of the genus Nephrotoma Meigen and of the
subgenus Tipula (Schummelia Edwards). In
Nephrotoma, a partly blackish sclerotized sp2 can be
found in for instance the Palaearctic aurantiocincta
Alexander, biarmigera Alexander, medioproducta
Alexander and the Oriental citricolor Alexander, inte-
gra Alexander, progne Alexander, subumbonis
Alexander, and umbonis Alexander. In the pertaining
species of Nephrotoma, the general shape of sp2 differs
from that of the species of Savtshenkia in that sp2 is
short and broad, anteriorly notched, and rather deep-
ly concave. All species of Schummelia studied (viz. ah-
rensi Savchenko, butzi Edwards, variicornis
Schummel, yerburi Edwards, zernyi Mannheims, and
zonaria Goetghebuer) have the anteromedial margin
of sp2 more or less darker sclerotized than the re-
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
mainder of the structure. The general shape of sp2 in
the species of Schummelia is similar to the elongate
form as found in the majority of species of
Savtshenkia. The phylogenetic positions of the per-
taining species of Nephrotoma and Schummelia with-
in the respective taxa are still unestablished. Except
for the similar modification of sp2, there are at the
moment, however, no indications to suggest that ei-
ther Nephrotoma or Schummelia, or both combined,
represent(s) the closest relative of Savtshenkia (see al-
so next character).
54. — Sp2: (0) dorsally separate from tergite 9-10;
(1) dorsally fused with ventromedial part of tergite
9-10.
Sp2 and tergite 9-10 are separate from each other
in the greater majority of Tipulidae, a situation con-
sidered the groundplan condition of the Lunatipula
group and Mediotipula. More than half of the species
of Savtshenkia also show this character state (fig. 100,
arrow). In the species of the clade fragilina to omega,
sp2 is fused with the ventromedial sclerotization of
tergite 9-10 (fig. 101). Both structures are firmly con-
nected over their entire width. Outside Savtshenkia, I
found a similar character state in species of the subge-
nus Zipula (Schummelia) only. In the examined spe-
cies of Schummelia (see previous character), sp2 and
the ventral sclerotization of tergite 9-10 are approxi-
mated. Some of these species show a partial fusion of
sp2 and tergite 9-10 (viz. ahrensi, variicornis, and yer-
burt). In zernyi, both structures are more firmly con-
nected. The studied species of Schummelia all have a
western Palaearctic distribution and represent only a
minor fraction of the subgenus which contains about
75 species distributed further in the Afrotropical,
Nearctic and especially Oriental Regions. Fusion of
tergite 9-10 with sp2 probably evolved independent-
ly in Schummelia. As stated under the preceding char-
acter, I do not consider Schummelia a potential sister
group of Savtshenkia.
55. — Sternite 8, posterior margin: (0) unmodified;
(1) caudally extended.
An unmodified sternite 8 in Tipulidae consists of a
relatively short structure of which the posterior mar-
gin is evenly convex or slightly concave and lacks any
trace of spinosity. This situation probably represents
the groundplan condition of this character for the
Lunatipula group and Mediotipula. In a number of
species of Savtshenkia, a modification of the posterior
margin of sternite 8 is found, which generally consists
of a posterior elongation of the structure (figs. 103,
KO MIO Aa WS LIS 173, 185 199, 90).
Elongation of sternite 8 is usually combined with the
presence of spines or strong setae along the posterior
margin (character 57). A caudally produced posterior
291
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 113-128. — 113-117, male sternite 8, posterior margin; 113, 7: (S.) cyrnosardensis, posterior view; 114, 115, 7. (S.) ha-
ennii, 114, posterior view; 115, ventral view; 116, 7. (S.) signata, posterior view; 117, 7. (S.) mannheimsi, posterior view; 118-
120, male sternite 9, anteroventral appendage, dorsal view; 118, 7: (S.) sordidipes, 119, T. (S.) eugeni; 120, T. (S.) limbata;
121-128, male tergite 9-10; 121, 122, 7. (S.) grisescens, 121, dorsal view; 122, posterior view; 123, 7. (S.) hartigiana, dorsal
view; 124, 7. (S.) atlas, dorsal view; 125, 7. (S.) goriziensis, dorsal view; 126, T: (S.) cheethami, dorsal view; 127, T. (S.) ig-
nobilis, dorsal view; 128, 7. (S.) subnodicornis, dorsal view.
margin of sternite 8 is found in the species /undbladi,
multipicta, rufina, and those of the clade alpium to
omega. Within the latter clade a short sternite 8 is
present in aspromontensis, gimmerthali, interserta,
pechlaneri, sordidipes, and venerabilis (see fig. 111).
56. — Sternite 8, posterior margin: (0) variously
292
shaped; (1) with V-shaped ventral carina.
Sternite 8 is present as an evenly curved sclerite in
the presumed groundplan condition of the
Lunatipula group and Mediotipula. Within
Savtshenkia a variety of forms of the posterior margin
of sternite 8 is found. The two subspecies rufina rufi-
na and rufina maderensis are distinguished from the
remainder of the subgenus by the presence of a mid-
ventral V-shaped carina in the posterior part of ster-
nite 8 (posterior view, fig. 104).
57. — Sternite 8, posterior margin: (0) unarmed; (1)
provided with strong setae or black spines.
In most Tipulidae, sternite 8 is unarmed, a situa-
tion considered to represent the groundplan state of
this character for the Zunatipula group and
Mediotipula. About half of the species of Savtshenkia
have the posterior margin of sternite 8 provided with
strong setae or short spines (figs. 105-109, 113-117,
177, 178, 189, 190). The number of spines or setae
and their location varies among the different species
of Savtshenkia. The species with an armed posterior
margin of sternite 8 include baltistanica, multipicta,
and most species of the clade alpium to omega.
Within the latter clade a number of species show an
unarmed sternite 8, viz. interserta, macaronesica, orna-
ta, and the species of the clade fragilina to invenusta
(see figs. 110-112).
58. — Sternite 8, posterior margin: (0) variously
shaped; (1) with medial tongue-like extension,
posteriorly concave.
As stated above, a number of species of Savtshenkia
have a caudally produced posterior margin of sternite
8. The species holoptera, obsoleta, and pagana are dis-
tinguished among these by the peculiarly shaped pos-
terior extension of sternite 8, which appears as a
tongue-like blade which is caudally more or less emar-
ginate (figs. 106, 107). In most specimens of pagana,
the caudal margin of the posterior extension is nearly
straight, in the species holoptera and obsoleta it is dis-
tinctly concave.
59. — Sternite 8, posterior margin: (0) unarmed or
armed with more than 10 spinous setae; (1) number
of spinous setae less, ranging from seven to one.
The number of strong setae or spines on the poste-
rior margin of sternite 8 in the species of Savtshenkia
which have this structure armed usually is (consider-
ably) more than 10 (figs. 105, 113-117, 177, 178,
189, 190). In the species obsoleta, there are about 13
to 15 spines on the posterior margin of sternite 8 (fig.
106). The species holoptera and pagana have less than
10 spines along the posterior margin of sternite 8.
Sternite 8 of pagana usually has about four to eight
spinous setae, in holoptera the number varies between
one and three (fig. 108; see Eiroa, 1987).
60. — Sternite 8, posterior margin: (0) medial
membranous area absent, small or, if large, without
rim of longish setae; (1) membranous area broadly
V-shaped, anteriorly extending, its margins set with
long slender setae.
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
As noticed above, the groundplan condition of ster-
nite 8 for the Lunatipula group and Mediotipula prob-
ably consists of an unmodified sclerite. The posterior
margin of the sternite in this situation is uniformly
sclerotized. In Savtshenkia, sternite 8 generally has no
membranous area at its posterior margin or the mem-
branous area is restricted to a narrow zone (figs. 102,
103. 1052102 Wil, wy, WS, USS 177 189) The
three species fragilina, limbata, and subvafra are distin-
guished from the remainder of Savtshenkia by a large
and V-shaped membranous area which is bordered by
a fringe of thickly set longish slender setae (fig. 110).
The species aster and confusa are the only other species
of Savtshenkia that also have a relatively large membra-
nous area on the posterior margin of sternite 8. In as-
ter, a V-shaped membranous area is present in
between the caudolateral extensions, the extensions
themselves are provided with a few black spines (fig.
109; see Appendix C). The species confusa has a broad
and rounded membranous zone (fig. 108; see
Appendix C). In both species, the margin of the mem-
branous area is surrounded by scattered setae only.
61. — Sternite 8, posterior margin: (0) ventrally
directed medial extension absent; (1) unarmed
medial extension present rostrally of lateral
extensions, or at this position with sclerotized edge.
In the majority of species of Savtshenkia, the poste-
rior margin of sternite 8 medially either is unmodified
or produced into a rather broad lobe provided with
spines or strong setae (figs. 102, 103, 105-111, 158).
An unarmed medial extension rostrally of a pair of
lateral extensions on the posterior margin of sternite 8
can be found in the three subspecies of invenusta and
in the species of the clade nielseni to omega. The size,
shape and sclerotization of the medial extension var-
ies among the species involved. In the subspecies of
invenusta, and in mannheimsi, omega, and trinacria
the medial extension is relatively small and moderate-
ly sclerotized (figs. 112, 117, 177, 178, 189, 190), it
is large and partly membranous in nielseni, whereas it
is more pronounced and heavily sclerotized in most
other species involved (figs. 114, 116). It should be
noted, however, that the species benesignata, boreosig-
nata, and eleonorae lack a protruding medial exten-
sion. In these three species, a sclerotized rim marks
the position of the medial extension.
62. — Sternite 8, posterior margin: (0) extension(s)
absent or dorsoventrally flat; (1) lateral extensions
appearing as bulbous structures, densely set with
spines.
The extensions of sternite 8 are relatively flat and
follow the curve of sternite 8 when seen in posterior
view in about half of the species of Savtshenkia with a
modified sternite 8. Most members of the clade cyr-
293
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
LAN
| ire va NUE
STZ |
Sn
Figs. 129-141. Male tergite 9-10.— 129, 7. (S.) baltistanica, dorsal view; 130, 7. (S.) elgonensis, posterior margin, dorsal view,
after celluloid preparation holotype; 131, 132, 7. (S.) aberdareica, 131, posterior view; 132, dorsal view; 133, T. (S.) sordidi-
pes, dorsal view; 134, 7. (S.) serrulifera, posterior view; 135, 7. (S.) aspromontensis, posterior view; 136, T. (S.) invenusta in-
venusta, dorsal view; 137, T. (S.) nielseni, dorsal view; 138, 7. (S.) boreosignata, dorsal view; 139, T. (S.) persignata tofina, pos-
terior view; 140, 7. (S.) signata, posterior view; 141, 7. (S.) mannheimsi, posterior view.
nosardensis to omega are characterized by the presence
of bulbously swollen lateral extensions of sternite 8.
In the majority of these species, the lateral extensions
are thickly set with black spines (figs. 113-116), but
in the species boreosignata only a few spines on the lat-
eral lobes are found. Moreover, the three species
mannheimsi, omega, and trinacria have a relatively flat
pair of lateral extensions (figs. 117, 178, 190; charac-
ter 65).
294
63. — Sternite 8, posterior margin: (0) lateral
extensions absent, or, if present, dorsally not
connected; (1) dorsally connected by a sclerotized
bridge.
In most species of Savtshenkia which carry a pair of
lateral extensions on the posterior margin of sternite
8, the extensions are dorsally separated by a membra-
nous area (figs. 113, 116, 117, 178, 190). The species
and subspecies corsosignata, haennii, subsignata subsig-
nata, and subsignata cazorla are distinguished by the
presence of a sclerotized dorsal bridge that connects
the lateral extensions of sternite 8 (fig. 114).
64. — Sternite 8, posterior margin: (0) variously
shaped; (1) in between medial and lateral extensions
a pair of medially directed projections.
The ventral margin of sternite 8 in between the
medial and lateral extensions is straight or evenly
curved in most species of Savtshenkia with a three-
lobed sternite 8. As an extra feature, corsosignata,
haennii, subsignata subsignata, and subsignata cazorla
have an acute and dorsomedially directed sclerotized
projection on each side of the medial extension (fig.
115, arrow).
65. — Sternite 8, posterior margin: (0) variously
shaped; (1) lateral extensions dorsoventrally
flattened, posterior margin straight, spines confined
to posterior margin.
The bulbously swollen lateral extensions of sternite
8 in most species of the clade cyrnosardensis to omega
are more or less circular in cross section and covered
with spines on their posterior half, dorsally as well as
ventrally (figs. 113-116). The species mannheimsi,
omega, and trinacria have the lateral extensions of
sternite 8 dorsoventrally flatter and straight along
their posterior margins (figs. 117, 177, 178, 189,
190). The spines in these three species are concentrat-
ed along the posterior margin of the sternite.
66. — Sternite 9, midventral anterior appendage: (0)
apically rounded; (1) apically bifid.
Sternite 9 often carries a more or less developed an-
terior midventral appendage in the Tipulidae. When
present, this appendage has a rounded anterior mar-
gin, as it has in the majority of species of Savtshenkia
(figs. 118, 120). In the species aspromontensis, aster,
eugeni, and serrulifera the anterior appendage has a bi-
fid apex (fig. 119).
67. — Sternite 9, midventral anterior appendage: (0)
a plain structure; (1) dorsally provided with medial
carina.
In the presumed groundplan condition of the
Lunatipula group and Mediotipula, the ventromedial
part of sternite 9, including the anterior appendage, is
a dorsoventrally flattened structure. The majority of
species of Savtshenkia show an anterior midventral
appendage of sternite 9 which looks like a simple
tongue-like extension without further modifications
(fig. 118). A different form of this structure is present
in the species of the clade fragilina to omega, where
the appendage is provided with a medial carina on its
dorsal side (fig. 120). Although the height of the cari-
na varies among the pertaining species, it is usually
H. pe Jone: Phylogeny of Tipula (Savtshenkia)
well developed and clearly visible. A similar anterior
appendage was also found in the species hartigiana.
Within the clade fragilina to omega only pechlaneri,
sardosignata, and subsignata cazorla lack a distinct me-
dial carina.
68. — Tergite 9-10, posterior margin: (0) extensions
variously shaped; (1) narrowing toward caudally
produced tip.
In Tipulidae, tergite 9-10 often has a pair of low
and rounded extensions at its posterior margin, a sit-
uation that probably represents the groundplan con-
dition of the Lunatipula group and Mediotipula.
Within Savtshenkia, only a few species show similar
extensions) (figsml2 15) 1225125) 126012746157)
The species concerned lie in the range alpha to ignob-
ilis, with the exception of the species atlas, brevianten-
nata, and hartigiana. In the three latter species, the
extensions are caudally produced and constricted be-
fore their tips. The medial margins of the extensions
in these species are slightly concave in dorsal view
(figs. 123, 124).
69. — Tergite 9-10, posterior margin: (0)
posteroventral sclerotization uninterrupted along
posterior margin; (1) separated into two parts by
medial incision.
Tergite 9-10 in Tipulidae often has a sclerotized
rim running along the ventral side of the posterior
margin. An uninterrupted rim probably represents
the groundplan state of this character for the
Lunatipula group and Mediotipula. The majority of
species of Savtshenkia also have a complete and broad
posteroventral sclerotized rim along the posterior
margin of tergite 9-10 (figs. 121, 123, 125-129, etc.,
after macerated specimens). Seen in dorsal view, this
rim seems to be intersected by the narrow and deep
medial incision of tergite 9-10 in the species atlas and
breviantennata (fig. 124).
70. — Tergite 9-10, posterior margin: (0) tips of
extensions broadly separate; (1) close together,
separated by a narrow gap.
A broad emargination or medial notch usually sep-
arates the extensions on the posterior margin of ter-
gite 9-10 in Tipulidae, a situation reflecting the pre-
sumed groundplan condition of the Lunatipula group
and Mediotipula. In the majority of species of
Savtshenkia, the posterior extensions of tergite 9-10
are separated by a broad V-shaped or U-shaped emar-
gination (figs. 121-125, 127, 129, etc.). In cheethami
and simulans, the extensions approach each other me-
dially and are separated by a rather deep and narrow
fissure (fig. 126).
71.—Tergite 9-10, posterior margin: (0) extensions
295
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
RCA
We
We
147
Figs. 142-151. — 142, 143, genital fork, dorsal view; 142, Tipula (Savtshenkia) hartigiana, 143, T. (S.) atlas, 144, 145, ovi-
positor, lateral view; 144, T° (S.) grisescens, 145, T° (S.) hartigiana, 146, 147, spermatheca; 146, 7. (S.) hartigiana; 147, T. (S.)
villeneuvit; 148, 149, female sternite 8 and hypogynial valves, ventral view; 148, 7. (S.) grisescens 149, T. (S.) ignobilis, 150,
151, female sternite 9, medial part, ventral view; 150, 7: (S.) jeekeli; 151, T. (S.) obsoleta.
posteriorly directed and rounded at tip; (1) laterally
curved and pointed at tip.
As noted above, the groundplan condition of ter-
gite 9-10 for the Lunatipula group and Mediotipula
probably shows a pair of broad and rounded exten-
sions on the posterior margin of the tergite. The ma-
jority of species of Savtshenkia, however, have the tips
of the extensions of tergite 9-10 pointed and out-
wardly curved when seen in dorsal view (figs. 128-
130, 137, 139-141, 174, 175, 186, 187).
72. — Tergite 9-10, posterior margin: (0) provided
with black spines; (1) spines absent.
The section of the family Tipulidae to which
Savtshenkia and its outgroups belong, is, among oth-
er things, characterized by the presence of black
spines on the posterior margin of tergite 9-10.
Although such spines are absent in the majority of
species of the Lunatipula group, there are several spe-
cies within the group which have the posterior mar-
gin of tergite 9-10 armed with spines. For this reason
I, putatively, consider this latter state the groundplan
296
condition of the Lunatipula group. All species of
Mediotipula and the majority of species of
Savtshenkia also show a spinose posterior margin of
tergite 9-10 (figs. 121-128, 133-136, 139, 140, 187).
Within Savtshenkia, total absence of spines on the
posterior margin of tergite 9-10 is found in the spe-
cies alpha (figs. 156, 157) and the species of the clade
baltistanica to chrysocephala (figs. 129-132). The spe-
cies glaucocinerea has a few knob-like protuberances
on the lower margin of the extensions of tergite 9-10,
but lacks distinct spines. Spines along the ventral
margin of the extensions of tergite 9-10 are absent in
the species mannheimsi and omega, but in both species
a few small spines are present in the dorsal part of the
medial cleft (figs. 141, 175).
73. Tergite 9-10, posterior margin: (0) extensions
variously shaped, usually long and laterally directed;
(1) extensions present as short spineless points.
The laterally directed extensions of the species of
the clade subnodicornis to omega usually terminate in
slender and elongate points (figs. 128-130, 137-141,
174, 175, 186, 187). A number of Afrotropical spe-
cies of Savtshenkia are characterized by the presence
of short and acute extensions, viz. the species aberda-
reica, akeleyi, asbolodes, chrysocephala, draconis, and
hancocki (figs. 131, 132).
74. — Tergite 9-10, posterior margin: (0) spines
confined to the extensions; (1) spines present above
medial notch.
As is usual in Tipulidae which have tergite 9-10
provided with spines, the spines are concentrated on
the posterior margin of the tergite in the majority of
species of Savtshenkia (figs. 121-128, 139, 140, 175,
187). Spines are present also in the area above the me-
dial notch of tergite 9-10 in the species aspromonte-
nsis, aster, eugent, odontostyla, serrulifera, sordidipes,
and venerabilis (figs. 133-135). In the latter five spe-
cies, there are about 10 spines in this area (figs. 133,
134), while aspromontensis and aster carry 20 to 30
spines above the notch (character 78).
75. — Tergite 9-10, posterior margin: (0) variously
shaped; (1) broadly V-shaped emarginate with
rounded corners.
The Kashmirian species sordidipes and venerabilis
differ from all other species of Savtshenkia by the
shape of the posterior margin of tergite 9-10. Tergite
9-10 in both species has a broad V-shaped emargina-
tion with a small median notch, while the posterior
corners are broadly rounded off (fig. 133). The spe-
cies sordidipes and venerabilis resemble each other very
much, including in details of the male terminalia.
Differences are found in the colour of the legs and ab-
domen and in the height of the anterior part of the in-
ner gonostylus.
76. — Tergite 9-10, posterior margin: (0) single pair
of extensions present; (1) two pairs present.
As in most other Tipulidae, the standard number
of extensions on the posterior margin of tergite 9-10
in Savtshenkia is two (figs. 121-127, 129-133, 136-
141, 156, 157, 174, 175, 186, 187). The species ser-
rulifera, aspromontensis, and aster differ from this by
having a medial and a sublateral pair of extensions.
Both pairs of extensions are provided with black
spines in aster and serrulifera (fig. 134), in aspro-
montensis the sublateral pair is unarmed (fig. 135).
77.—Tergite 9-10, posterior margin: (0) dorsal
surface of extensions rounded; (1) flattened.
In general, the extensions on the posterior margin
of tergite 9-10 are rounded on their dorsal surface in
Tipulidae, a situation found also in the majority of
species of Savtshenkia. As a special trait, the species as-
promontensis and aster have the medial pair of exten-
sions of the posterior margin of tergite 10 flattened.
H. DE Jone: Phylogeny of Tipula (Savtshenkia)
The dorsal surface of the medial extensions of both
species is glossy black.
78. Tergite 9-10, posterior margin: (0) dorsal
surface in between extensions unarmed or armed
with up to 10 spines; (1) dorsal surface in between
extensions armed with 20 to 30 spines.
As discussed under character 74, a few species of
Savtshenkia have the area above the medial notch of
tergite 9-10 armed with up to 10 spines (fig. 133,
134). The number of spines is higher in the species
aspromontensis and aster. In aspromontensis, there are
about 30 spines above the medial notch (fig. 135), in
aster there are about 20.
79. — Tergite 9-10, posterior margin: (0) extensions
present; (1) margin straightened, with small medial
notch.
As discussed above, the majority of species of
Savtshenkia have a pair of lateral extensions on the
posterior margin of tergite 9-10, either low and
rounded or elongate, diverging and pointed (figs.
128-130, 134, 135, 137-141, 174, 175, 186, 187).
The species interserta, the three subspecies of invenus-
ta, and the species subvafra are characterized by a
straightened posterior margin of tergite 10 (fig. 136).
In subvafra, a pair of small posteroventral projections
flank the narrow medial notch; such projections are
absent in interserta and invenusta.
80. — Tergite 9-10: (0) broader than long, or about
as long as broad; (1) elongate, distinctly longer than
broad.
In Tipulidae, tergite 9-10 usually is a relatively
short and broad structure, a situation which probably
reflects the groundplan condition of the Lunatipula
group and Mediotipula. The majority of species of
Savtshenkia also show a similar character state (figs.
121, 123-129, 132, 133, 136, 156). In the pertaining
species, tergite 9-10 is broader than long and in most
of the species shows a broad medial membranous
zone. The species and subspecies of the clade nielseni
to omega differ from this in having an elongate tergite
9-10 which is markedly longer than broad (fig. 137,
138, 174, 186). The medial membranous area of ter-
gite 9-10 in these taxa is restricted to a narrow zone,
with the exception of the species boreosignata which
has a relatively wide membranous area (fig. 138).
81. Tergite 9-10, posterior margin: (0) extensions
variously shaped; (1) rather broad, ventral margin
straightened, terminating in an acute lateral tip.
As noted above (character 71), most species of
Savtshenkia have the extensions on the posterior mar-
gin of tergite 9-10 laterally directed and pointed at
the tip. In the majority of these species, the ventrome-
297
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
dial margin of the extensions is distinctly concave
when seen in posterior view (fig. 139). The species
corsosignata, mannheimsi, omega, signata, and trina-
cria are characterized by rather broad extensions of
tergite 9-10 of which the ventral margin appears to be
almost straight in posterior view (figs. 140, 141, 175,
187).
Female terminalia
82. — Genital fork: (0) broadest width in posterior
half; (1) broadest width in anterior half.
In general, the genital fork in Tipulidae is a rather
slender structure that widens in its apical posterior
part. This situation probably represents the ground-
plan condition of the Lunatipula group. Within
Mediotipula, the genital fork is slender throughout,
with the widest portion lying in the anterior part. The
general structure of the genital fork is rather uniform
in the species of Savtshenkia, being broad with its
greatest width lying in the anterior half (figs. 142,
169, 201; lower side in illustrations). The species at
las and breviantennata are the only studied represen-
tatives of Savtshenkia which have the broadest part of
the genital fork situated in the posterior section (fig.
143; upper side in illustration).
83. — Hypogynial valve, dorsal margin: (0) not
blackish sclerotized; (1) blackish sclerotized.
The hypogynial valves are usually uniformly sclero-
tized in Tipulidae and this condition probably repre-
sents the groundplan state of this character for the
Lunatipula group and Mediotipula. All species of
Savtshenkia of which the female was examined have
the dorsal margin of the hypogynial valve blackish
sclerotized.
84. — Ovipositor: (0) well developed, elongate; (1)
short.
The ovipositor in Tipulidae generally is well devel-
oped and has elongate cerci and hypogynial valves, a
situation which represents the groundplan condition
of the Lunatipula group and Mediotipula. The same
state occurs in the majority of species of Savtshenkia
(figs. 145, 165, 197). Cerci and hypogynial valves are
short in the females of grisescens, holoptera, obsoleta,
pagana, and subnodicornis (fig. 144).
85. — Spermathecae: (0) globular; (1) elongate.
Theowald’s (1957, 1967) arrangement of species groups of Tipula (Savtshenkia) based on larvae and pupae (1957), and pu-
Table 2
pae (1967).
1957
rufina group cheethami
rufina
subnodicornis group goriziensis
(not mentioned)
subnodicornis
subvafra group atlas (as breviantennata)
signata group alpium
confusa
nielseni
obsoleta
(not mentioned)
(not mentioned)
signata
staegeri
Table 3
1967
cheethami
rufina
(not filed)
grisescens
subnodicornis
atlas (as breviantennata)
alpium
confusa
(not mentioned)
obsoleta
odontostyla
pagana
signata
staegeri
Division of the Palaearctic species and subspecies of Tipula (Savtshenkia) into species groups according to Savchenko (1961)
and Theowald (1973, 1978). Theowald’s group numbering is given in parentheses, a ‘?’ denotes a species placed by Theowald
under restriction in the pertaining group.
Notes: 1: placed in miscellaneous group by Savchenko, under restriction in convexifrons group by Theowald; 2: referred to by
Theowald only; 3: placed in rufina group by Savchenko, in pagana group by Theowald; 4: placed in miscellaneous group by
Savchenko, under restriction in pagana group by Theowald; 5: placed in cheethami group by Savchenko, in alpium group by
Theowald; 6: placed in rufina group by Savchenko, under restriction in obsoleta group by Theowald; 7: referred to by
Savchenko only. ni
298
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
Savchenko 1961 Theowald 1973, 1978 Species and subspecies
subnodicornis group subnodicornis group (1) postposita
subnodicornis
convexifrons group (2) convexifrons
interserta
? venerabilis È
limbata group grisescens group (3) goriziensis°
grisescens (as macrocera in Savchenko 1961)
villeneuvii (as goriziensis villeneuvit)
invenusta group (4) invenusta invenusta
invenusta subinvenusta
subvafra
limbata group (5) limbata
pagana group (6) baltistanica’
glaucocinerea
holoptera
opter (as pagana holoptera)
? kiushiuensis
? koreana*
B 2
? minuscula” (as nana)
nivalis
pagana (as pagana pagana)
gimmerthali group (7) gimmerthali
pechlaneri”
rufina group rufina group (8) rufina rufina
rufina maderensis
alpium group (11) alpium
subalpium
macaronesica”
baltistanica *
odontostyla °
cheethami group atlas group (9) ae i
reviantennata
cheethami
macaronesica °
marmorata group obsoleta group (10) aspromontensis”
lundbladi
? mohriana
multipicta È
obsoleta
? odontostyla°
serrulifera
marmorata group (12) confusa (as marmorata)
eugeni
jeekeli
staegeri group (13) boreosignata 7
nielseni’
sardosignata
staegeri
subsignata
signata group (14) benesignata
mannbeimsi *
persignata persignata
persignata tofina
signata
miscellaneous group koreana”
letifera’
venerabilis È
299
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
The shape of the (three) spermathecae in Tipulidae
is usually spherical to somewhat ovoid. The ground-
plan condition of this character for the Lunatipula
group and Mediotipula is probably represented by this
state. The majority of females of Savtshenkia exam-
ined also show this spermathecal form (figs. 146, 170,
202). Only the species cheethami, goriziensis, simu-
lans, and villeneuvii are characterized by the presence
of elongate spermathecae (fig. 147). It should be not-
ed that the female of tu/ipa is not yet known.
86. — Sternite 8: (0) ventromedial membranous area
near base of hypogynial valves V-shaped,
encompassing base of medial margins of hypogynial
valves; (1) posteriorly terminating in between
hypogynial valves.
The ventromedial membranous area of sternite 8
broadens posteriorly and encompasses the bases of the
hypogynial valves medially in the species of the
Lunatipula group and Mediotipula, thus probably
representing the groundplan condition of this charac-
ter for the pertaining taxa. The same state is found in
the species of Savtshenkia which lie in the range alpha
to simulans (fig. 148). The membranous area lies in
between the bases of the hypogynial valves in the spe-
cies of the clade ignobilis to omega (fig. 149). In these
species, the valves are usually broadly separate at the
posterior end of the membrane.
87. — Sternite 9: (0) with long and slender medial
projection; (1) with broad and short medial
projection.
The medial projection of sternite 9 usually takes
the form of an elongate slender and pointed structure
in Tipulidae. This situation, which probably reflects
the groundplan condition of the Lunatipula group
and Mediotipula, is present in the majority of species
of Savtshenkia (figs. 150, 168, 200). The species ben-
esignata, eugeni, holoptera, obsoleta, and pagana have a
differently built medial projection, which consists of
a rather broad structure that is narrowed near mid-
length (fig. 151). In eugeni, the cavities next to the
medial projection are restricted to the anterior part of
sternite 9, whereas they run along the whole length of
the medial projection in the four other species.
DISCUSSION OF ADOPTED PHYLOGENY
The character state matrix as given in table 4 was
used to establish the phylogeny of the species and
subspecies of Savtshenkia. The character states are bi-
nary coded, polymorphies are coded —’, absent data
Initially all characters employed had the same
weight 1. The character state matrix of table 4 was
run under the command ‘mh*; bb*;’ of the parsimo-
ny program Hennic86. The first part of the com-
300
mand (mh*;) constructs several trees to which it ap-
plies branch-swapping, retaining one tree for each ini-
tial tree, while the second part of the command (bb*;)
applies extended branch-swapping to each of the in-
put trees, retaining all most parsimonious trees.
Although it does not guarantee to find all trees of mi-
nimal length, applying ‘mh*; bb*; to extensive data
sets offers a good alternative for the time-consuming
implicit enumeration options of HENNIG86 that de-
finitely result in finding all trees of minimal length
(Farris 1988). The procedure followed resulted in 28
equally most parsimonious trees with length 157,
consistency index 55, and retention index 88.
Subsequently successive weighting was carried out
under the command ‘mh*; bb*; xs w; cc;’. This proce-
dure uses the fit of the characters to the input trees to
calculate the weights as the product of the character
consistency and character retention indices, which are
then scaled in the range 0-10 (Farris 1988).
Successive weighting resulted in 14 equally most par-
simonious trees with length 723, consistency index
84, and retention index 96. This result was obtained
after a single weighting run and did not improve
upon further repeats. The calculated weights for the
characters are given in table 5, the strict consensus
tree of the 14 equally most parsimonious trees is
shown in fig. 2. The strict consensus tree of the 14
trees resulting of successive weighting (fig. 2) differs
from the strict consensus tree of the 28 trees without
successive weighting in the presence of the clade bal
tistanica to chrysocephala in the former. The internode
leading to this clade is not recognized in the consen-
sus tree of the 28 trees without weighting, lowering
the grouping baltistanica to chrysocephala one level
relative to its position in fig. 2. For the remainder the
topologies of both consensus trees are identical. The
results of the phylogenetic analysis will be discussed
with reference to fig. 2.
At the base of the cladogram a trichotomy of the
clades alpha and grisescens, hartigiana to simulans, and
ignobilis to omega is found. The species pair alpha and
grisescens probably are sister species as exemplified by
two homoplasies and one reversal [characters 2
(length of male antenna), 9 (size of gonapophysis),
and 4 (shape of cell m1), respectively].
The second clade seems to be better substantiated
by the characteristic shape of the gonapophysis (char-
acter 10) and the acute posterior corner of the base of
the inner gonostylus (character 32). The first lineage
within this clade leads to a triplet containing the spe-
cies hartigiana as the sister species of the pair atlas and
breviantennata. In its original description, hartigiana
was referred to the atlas group sensu Theowald (table
3) (Theowald, Dufour & Oosterbroek 1982). Besides
the species atlas, breviantennata, and hartigiana, the
atlas group sensu Theowald also included the species
cheethami. As the present analysis shows, the latter
species seems to be more closely related to the species
of the second group within the clade hartigiana to
simulans. This group, the clade tulipa to simulans,
consists of the species tulipa, villeneuvii, goriziensis,
and the pair cheethami and simulans. Dufour (1983),
on describing the species tulipa, placed it in the grises-
cens group sensu Theowald which by then contained
the species goriziensis, grisescens, and villeneuvii (table
3). Dufour (1983) suggested that grisescens stands
apart from the other three species, including villeneu-
vii, thus countering Theowald & Oosterbroek’s
(1981) contention that vi/leneuvii should be regarded
a subspecies of grisescens. The results of the present
analysis endorse Dufour’s views.
The third basal clade in fig. 2 contains the remain-
der of Savtshenkia. It opens with the successive
branching off of the species ignobilis and subnodicor-
nis, followed by the remainder of species of
Savtshenkia which constitute the monophyletic group
baltistanica to omega. The species subnodicornis was
placed either near goriziensis, grisescens, or postposita
by Savchenko (1961) and Theowald (1957, 1967,
1973, 1978) (tables 2, 3). The species listed belong to
the phylogenetically more basal species of
Savtshenkia, but none of them appears to be the actu-
al sister species of subnodicornis.
The clade baltistanica to chrysocephala contains the
majority of species of Savtshenkia with a non-spinous
posterior margin of tergite 9-10 (character 72). The
only other species of Savtshenkia in which the spines
on the extensions of tergite 9-10 are absent, are the
phylogenetically basal species alpha and the derived
species mannheimsi and omega. The basal trichotomy
of the clade baltistanica to chrysocephala is constituted
of the species baltistanica, the species pair glaucocine-
rea and postposita, and the clade lundbladi to chrysoce-
phala. The clade lundbladi to chrysocephala contains
almost all strictly Atlantic (viz. /undbladi, multipicta,
and rufina maderensis), and all Afrotropical species of
Savtshenkia (viz. nephrotomoides to chrysocephala).
(The only remaining Atlantic species of Savtshenkia is
macaronesica from the Azores, a species closely related
to alpium, see below.) As table 3 shows, the
Palaearctic species of the clade baltistanica to chrysoc-
ephala were placed in various groupings by
Savchenko (1961) and Theowald (1973). (The phy-
logeny of the Afrotropical species presented here
should be regarded as preliminary only. Most species
are known of just a few specimens, thus giving the
strong impression that the Afrotropical fauna is poor-
ly known. Furthermore, the species are in general very
similar to each other and homoplasy in this group
seems to be rather common. As the present study is
primarily concerned with the western Mediterranean
species of Savtshenkia, this issue is not further ex-
H. DE Jone: Phylogeny of Tipula (Savtshenkia)
plored here.)
Modifications of the posterior margin of the male
sternite 8 (characters 55, 57) suggest that the remain-
der of species of Savtshenkia, the clade alpium to ome-
ga, constitute a monophyletic group. The contours of
this group were recognized by Theowald (1957,
1967) in his signata group (table 2), by Savchenko
(1961) in his marmorata group, and by Theowald
(1973) in his obsoleta, alpium, marmorata, staegeri,
and signata groups (table 3). Within the clade alpinum
to omega, the basal lineage is composed of the species
pair alpium and macaronesica. Theowald (1973) cor-
rectly grouped both species in his alpium group, but
Savchenko (1961) placed macaronesica in his cheetha-
mi group (table 3). The species subalpium is probably
closely related to alpium and macaronesica (see
Appendix B). The sister group of alpium and macar-
onesica is substantiated by apomorphies of the sperm
pump (characters 48, 49). The basal phylogeny of
this clade (jeekeli to omega) could not be fully re-
solved. Four lineages constitute a polytomy, with the
first containing the single species jeekeli, the second
the clade obsoleta to pagana, the third the clade confu-
sa to aster, and the fourth the clade fragilis to omega.
The species of the clade obsoleta to pagana are most
readily distinguished from the other species of
Savtshenkia by the peculiar form of sternite 8 (charac-
ter 58). Notwithstanding this, Savchenko (1961) as
well as Theowald (1973) placed obsoleta in a species
group different from holoptera and pagana (table 3).
[I here retain holoptera as a taxon separate from paga-
na. Both taxa are usually distinguished by the length
of the wing and build of the legs in the female (mac-
ropterous with slender legs in holoptera, brachypte-
rous with stoutish legs in pagana), and the shape of
the posterior margin of male sternite 8 and the num-
ber of setae it carries (more deeply emarginate with
one seta on tubercle in holoptera, less emarginate to
truncate with about five to seven setae on tubercles in
pagana) (Coe 1950; Edwards 1939). However, num-
bers of setae on the posterior margin of sternite 8
intermediate to those of typical holoptera and pagana
can be found in certain specimens (Eiroa 1987; see al-
so Hutson & Vane-Wright 1969). The occurrence of
intermediate stages of fully winged and brachypterous
forms has been recorded within Savtshenkia for the fe-
male of gimmerthali (Dufour & Brunhes 1984). As
long as a similar variability of wing length is not re-
corded for females of the pair holoptera and pagana, I
prefer to keep them apart. This the more so as the
studied male specimens of holoptera show a slight dif-
ference with those of pagana in the presence of a small
acute and blackish sclerotized extension at the poste-
rior end of the dorsal edge of the inner gonostylus. }
The clade confusa to aster, which is substantiated by
the elongate aedeagus (character 15) and the presence
301
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Table 4. Character state matrix of Tipula (Lunatipula), T. (Mediotipula), and the species and subspecies of 7. (Savtshenkia).
Character 1 2 3
0 0 0
Lunatipula 00000 00000 00000 00000 00000 00000 00000
Mediotipula = 00010 00000 00000 00000 00010 00000 00000
aberdareica 01010 00000 10000 00000 00001 00000 00000
akeleyi 01010 10000 10000 00001 00001 01000 00000
alpha 01000 00010 00000 00000 00000 00000 00000
alpium 00010 00010 00000 00000 10000 00000 00000
asbolodes 01010 00000 10000 00000 00001 00000 00000
aspromontensis 00010 00000 00001 000-0 00000 00110 00000
aster 00010 00000 00001 00000 00000 00110 00000
atlas 10010 01001 00000 00010 00000 00000 01000
baltistanica 00010 00000 00000 00000 00000 00000 00000
benesignata 00010 00000 00001 — 00000 01100 00000 00000
boreosignata 00010 00010 00000 00000 01000 00000 00000
breviantennata 10010 01001 00000 00010 00000 00000 01000
caligo 01010 10000 10000 00001 00001 01000 00000
cheethami 00011 00001 00000 00010 00100 00000 01100
chrysocephala 01010 10000 10000 00001 00001 01000 00000
confusa 00010 00000 00001 00000 00000 00000 00001
corsosignata 00010 00100 00101 00100 01010 10001 00000
cyrnosardensis 10010 00000 01001 00000 00010 00000 00000
draconis 01010 00000 00000 00000 00001 00000 00000
eleonorae 00010 00000 00000 00000 01000 00000 00000
elgonensis 01010 00000 00000 00000 0000? 00000 00000
eugeni 00010 00000 00001 00000 00000 00000 00001
fragilina 00010 00000 00001 01000 00010 00001 00000
fragilis 00010 00000 01001 00000 00000 10000 00000
gimmerthali 00010 00000 00001 00000 00010 00001 00000
g. mattheyi 00010 00000 00001 00000 00010 00001 00000
Va pteromaculata 00010 00000 00001 00000 00010 00001 00000
glaucocinerea 00000 00000 00000 00000 00000 00000 00000
goriziensis 00010 00001 00000 00000 00100 00000 01100
graciae 00010 00000 00001 00000 00000 00000 00000
grisescens 01000 00010 00000 00000 00000 00000 00000
haennii 00010 00000 00101 00100 01010 10000 00000
hancocki 00010 00000 10000 00000 00001 00000 00000
hartigiana 00010 00001 00000 00010 00000 00000 01000
holoptera 01010 00000 00000 10000 00000 00000 00000
ignobilis 00010 00000 00000 00000 00000 00000 00000
interserta 00010 00010 00001 00000 00000 00000 00000
invenusta 00010 00000 00001 00000 00010 00000 00000
1. microinvenusta 00010 00000 00001 00000 00010 00000 00000
i. subinvenusta 00010 00000 00001 00000 00010 00000 00000
jeekeli 00010 00000 00000 00000 00000 00000 00001
limbata 00010 00000 00001 01000 00000 00001 00000
lundbladi 00110 00000 00001 00000 00001 00000 00000
macaronesica 00010 00010 00000 00000 10000 00000 00000
mannheimsi 00010 00000 00001 00000 01100 00000 10000
302
un
00000
00000
01000
01000
00000
01000
01000
01101
01111
00000
01000
01110
01010
00000
01000
00000
01000
01111
01110
01110
01000
01010
01000
01101
01110
01110
01110
01110
01110
01000
00000
01111
00000
01110
01000
00000
01110
00000
01101
01110
01110
01110
01110
01110
01000
01000
01110
00000
00000
00100
00100
00100
00101
00100
00100
00101
00100
00100
00111
01111
00100
00100
00100
00100
10101
00111
00111
00100
01111
00100
00101
00111
00101
00110
00110
00110
00100
00100
00101
00100
00111
00100
00100
00101
00100
10100
00111
00111
00111
00101
00111
00101
00101
00111
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
11000
11000
00000
00000
00000
00000
00000
11110
01000
00000
11000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
11110
00000
00000
00000
00000
00000
10000
10000
10000
00000
00000
00000
00000
11001
om
00000
00000
00000
00000
00000
00000
00000
10000
10000
00110
00000
01000
01000
00110
00000
00001
00000
00000
01000
01000
00000
01000
00000
10000
01000
00000
01000
01000
01000
00000
00000
00000
00000
01000
00000
01100
00000
00000
00000
01000
01000
01000
00000
01000
00000
00000
01000
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
00000
00000
11100
11100
01000
10000
11100
00010
00010
00000
11000
10000
00000
00000
11000
00000
11100
10000
10000
10000
11100
10000
11000
10010
10000
10000
10000
10000
10000
11000
00000
00000
00000
10000
11100
00000
10000
00000
00000
00000
00000
10000
10000
10000
11000
10000
11000
303
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Table 4. (continued) Character state matrix of Tipula (Lunatipula), T. (Mediotipula), and the species and subspecies of 7: (Savtshe |
Character 1 2 3
0 0
multipicta 00110 00000 00001 00000 00001 00000 00000
nephrotomoides 00010 00000 00000 00001 00001 00000 00000
nielseni 00010 00000 00001 00000 00010 00001 00000
obsoleta 00010 01000 00001 10000 00000 00000 00000
odontostyla 00010 00000 00001 00000 00010 00000 00000
omega 00010 00000 00001 00000 01100 00000 10000
ornata 00010 00000 00001 00000 00000 10000 00000
pagana 01010 00000 00000 10000 00000 00000 00000
pechlaneri 00010 00000 00001 00000 01010 00001 00000
persignata 00010 00000 00001 00000 01100 00000 10000
p. tofina 00010 00000 00001 00000 01100 00000 10000
phoroctenia 00010 00000 01001 00000 00000 00000 00000
postposita 00000 00000 00000 00000 00000 00000 00000
rufina 00010 00000 00000 00000 00001 00000 00000
r. maderensis 00010 00000 00001 00000 00001 00000 00000
sardosignata 00010 00000 00001 00000 00010 00000 00000
serrulifera 00010 00000 00001 00000 00000 00110 00000
signata 00010 00000 00001 00000 01100 00000 10000
simulans 00011 00001 00000 00010 00100 00000 01100
sordidipes 00010 00000 00001 00000 00000 00000 00000
staegeri 00010 00000 00001 00000 00010 00000 00000
subnodicornis 000-0 00000 00000 00000 00000 00000 00000
subsignata 00010 00100 00111 00100 01010 10000 00010
s. cazorla 00010 00100 00111 00100 01010 10000 00010
subvafra 00010 00000 00001 01000 00010 00001 00000
trinacria 00010 00000 00001 00000 01100 00000 10000
tulipa 00010 00001 00000 00000 00100 00000 01000
venerabilis 00010 00000 00001 00000 00000 00000 00000
villeneuvii 10010 00001 00000 00000 00100 00000 01000
of elongate and anteriorly directed posterior apo-
demes of the sperm pump (character 50), contains as
its first lineage the triplet confusa, interserta, and orna-
ta. Although differing in most genital features, these
three species share the presence of the quite remark-
able slender elongate and anteriorly extended posteri-
or apodemes of the sperm pump (character 51). In
fig. 2, the remainder of the clade confusa to aster is de-
picted as a monophyletic group which is substantiat-
ed by a single reversal only. The reversal concerns the
redirection and reduction of the extensions of the
posterior margin of tergite 9-10 (character 71). The
extensions are rather differently shaped in the species
of this grouping and its actual monophyly is open to
question. The species eugeni, which is included in this
grouping, has the extensions of tergite 9-10 well de-
veloped and laterally extended.
Theischinger (1983) correctly considered his spe-
cies aster to be the closest relative of aspromontensis.
Theowald (1973) placed the species confusa (as
304
marmorata), eugeni, and jeekeli in his marmorata
group on account of the peculiar shape of their inner
gonostyli (character 35). When other characters are
taken into consideration, the three species appear not
to be closely related, forcing the conclusion that the
characteristic shape of the inner gonostylus must be
due to homoplasy.
Next in fig. 2 is the clade fragilis to omega, in which
the Nearctic species pair fragilis and phoroctenia
branch off first, followed by the species odontostyla.
The remainder of species of Savtshenkia constitute
the monophyletic group fragilina to omega of which
the first lineage leads to the clade fragilina to invenus-
ta microinvenusta. This latter clade contains species
which are characterized by the presumably secondary
absence of spines on the posterior margin of male
sternite 8 (character 57). It is composed of species
which Theowald (1973) arrayed in his gimmerthali,
invenusta, and limbata groups (table 3), together with
the Nearctic species fragilina (the species mohriana
H. DE Jone: Phylogeny of Tipula (Savtshenkia)
5 6 7 8
0 0 0 0
01000 00101 01000 00000 00000 11000 00000 01100 10
01000 00100 00000 00000 00000 11000 00000 01100 10
11110 00111 01000 11000 01000 10000 00001 01100 10
01110 00101 01100 00000 00000 10000 00000 01110 11
01110 00101 01000 00000 00000 10010 00000 Ope eg Gs
01110 00111 01000 11001 01000 11000 00001 1988.89 BR
01111 10101 00000 00000 00000 10000 00000 01100 10
01110 00101 01110 00000 00000 10000 00000 01110 11
01110 00110 00000 00000 00000 10000 00000 01100 10
01110 00111 01000 11000 01000 10000 00001 ERS DE
01110 00111 01000 11000 01000 10000 00001 One Big 58
01110 00101 01000 00000 00000 10000 00000 01100 10
01000 00100 00000 00000 00000 11000 00000 0902 2?
01000 00101 10000 00000 00000 11000 00000 01100 10
01000 00101 10000 00000 00000 11000 00000 01100 10
11110 00111 01000 11000 00000 10000 00001 01100 10
01111 00101 01000 00000 10000 00010 10000 01100 10
01110 00111 01000 11000 01000 10000 00001 11100 10
00000 00100 00000 00000 00001 00000 00000 01101 00
01111 00100 01000 00000 00000 00011 00000 01100 10
01110 00111 01000 01000 01000 10000 00000 01100 10
00000 00100 00000 00000 00000 10000 00000 01110 10
01110 00111 01000 11110 01000 10000 00001 01100 10
01110 00111 01000 11110 00000 10000 00001 One? DE
01110 00111 00001 00000 01000 10000 00010 01100 10
01110 00111 01000 11001 01000 10000 00001 11100 10
00000 00100 00000 00000 00000 00000 00000 022723 DE
01111 00100 01000 00000 00000 00011 00000 Op ees 8
00000 00100 00000 00000 00000 00000 00000 01101 00
and productella probably also belong here; see
Appendix B).
The clade fragilina to invenusta microinvenusta
contains two trichotomies. The exact relationships of
the species fragilina, limbata and subvafra as well as
those between the three subspecies of gimmerthali are
hard to establish. The subspecies gimmerthali pterom-
aculata was described as differing from the nomino-
typical subspecies by its darker colour, spotted wings,
and relatively short second flagellomere (Theowald,
Dufour & Oosterbroek 1982). Examination of mate-
rial of the nominotypical form of gimmerthali, how-
ever, revealed that the length of the second flagel-
lomere varies considerably within this subspecies.
The relative length of the second flagellomere thus of-
fers no arguments to distinguish gimmerthali pterom-
aculata from the nominotypical subspecies. [The
length of the second male flagellomere has also been
used to distinguish gimmerthali from pechlaneri
(Mannheims & Pechlaner 1963). Considering the
variability of this character within gimmerthali it has
proved to be non-applicable here. The species gim-
merthali and pechlaneri are more easily distinguished
on account of the structure of the inner gonostylus
and the shape of male tergite 9-10 (see also Appendix
C). The monophyly of the three subspecies of gim-
merthali could be demonstrated by the shape of the
inner gonostylus only (character 41)]. Useful differ-
ences between gimmerthali gimmerthali and gimmer-
thali pteromaculata are found in the autapomorphous
larger outer gonostylus and relatively plump anterior
tip of the inner gonostylus of gimmerthali pteromacu-
lata (cf. figs. 66 and 67, and 83 and 84). The subspe-
cies gimmerthali mattheyi, known of the male holo-
type only (not examined), apparently differs from the
other subspecies by the autapomorphous absence of a
posterior projection on the outer gonostylus and the
reduced armature of the posterior margin of tergite 9-
10 (Theowald & Dufour 1983). [The females of both
gimmerthali mattheyi and gimmerthali pteromaculata
305
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Table 5. Weights of characters after single weighting run using the command ‘mh*; bb*; xs w; cc;’ of Hennig86.
Character Weight Character
1 1 31
2 1 32
3 10 33
4 1 34
5 10 35
6 10 36
7 2 37
8 10 38
9 1 39
10 10 40
11 10 41
12 2 42
13 10 43
14 10 44
15 1 45
16 10 46
17 10 47
18 10 48
19 3 49
20 3 50
21 10 51
22 4 52
23 4 53
24 2 54
25 10 55
26 2 56
27 10 57
28 10 58
29 10 59
30 2 60
Weight Character Weight
10 61 4
10 62 10
10 63 10
10 64 10
0 65 10
4 66 10
10 67 1
10 68 10
10 69 10
10 70 10
10 Vik 1
10 72) 2
10 73 4
2 74 4
2. 75) 10
10 76 10
10 ij 10
4 78 10
2 79 1
10 80 10
10 81 3
10 82 2
10 83 10
10 84 1
1 85 10
10 86 10
1 87 1
10
10
10
remain unknown. Those of gimmerthali gimmerthali
are usually brachypterous, but populations of the no-
minotypical form of gimmerthali are known from
Switzerland which have female wing lengths ranging
from the brachypterous to the fully winged condition
(Dufour & Brunhes 1984). Dufour (1990) further-
more recorded hemipterous females of gimmerthali
from the eastern side of the Sierra Nevada, Spain.]
The species invenusta presently contains three sub-
species. Differences between the subspecies can be
found in details of the male terminalia and in the ven-
tral distance between the eyes. The examined speci-
mens of invenusta invenusta conform to Riedel’s
(1919) description of the species as having rounded
posterolateral corners on tergite 9-10 [‘Lam. term.
sup. (...) mit abgerundeten Ecken’). However, illus-
trations of the terminalia of ‘invenusta invenusta by
Savchenko (1961) and Theowald (1973) depict ter-
gite 9-10 with acute posterior extensions.
Examination of material present in ZMAN showed that
Theowald actually figured a specimen that can only
be identified as invenusta subinvenusta. However,
considering Savchenko’s illustration there still is a
possibility that the structure of the posterior margin
of tergite 9-10 varies within invenusta invenusta. [All
306
three described subspecies bear spines on the postero-
ventral margin of tergite 9-10. Having examined sev-
eral of its paratypes, I can not corroborate Dufour’s
(1990) observation that spines are absent from the
posterior margin of tergite 9-10 in his invenusta mi-
croinvenusta.| Another complicating factor in main-
taining the three recognized subspecies is the observa-
tion that Tipulidae occurring at high geographical
latitudes or high altitudes usually have smaller eyes
than their relatives living under more moderate eco-
logical conditions. Therefore, the size of the eyes in
specimens of invenusta might well be an ecotypic dif-
ferentiation in response to local environmental condi-
tions and thus would be of little value in separating
the subspecies. The complicated nature of the taxon-
omy of invenusta is furthermore illustrated by the fact
that Alexander described invenusta invenusta under
three different names, viz. docilis (correctly synonym-
ized with invenusta by Savchenko 1961), and insig-
nifica and packardi (synonymized in this paper; see
Appendix A, synonymy). Hancock (1988) suggested
that yet another subspecies might be introduced to
accommodate a morphologically slightly deviating
population from Andorra (Pyrenees). I here would
like to stress the need to study the variability of this
taxon over its immense Holarctic range before even
considering to introduce new formal names for single
isolated populations. [The most recently published
map showing the Palaearctic distribution of invenusta
is the one given by Dufour (1992).]
The clade cyrnosardensis to omega contains the spe-
cies of Theowald’s (1973) staegeri and signata species
groups (table 3). Most species of this clade are easily
recognized by the presence of the bulbous and spi-
nous extensions on the posterior margin of sternite 8
(character 62). The clade opens with a trichotomy
consisting of the species cyrnosardensis and staegeri in
combination with a clade which includes the remain-
der of the species. Both cyrnosardensis and staegeri are
species distinguished by a number of autapomorphies
but apparently lack any further character states that
could enable the resolution of their exact phylogenet-
ic position.
The species nielseni and sardosignata are depicted
here as sister species as is suggested by their broad
based outer gonostyli, although the overall shape of
this structure differs remarkably in both species (char-
acter 46). The two species are, moreover, distin-
guished from each other by the autapomorphous
form of the inner gonostylus and the shape of sternite
8. Their tentative sister species relationship could not
be substantiated by other apomorphies.
Although on quite different grounds, the clade
haennii to subsignata cazorla contains the four taxa
which were recognized as a monophyletic group by
Dufour (1991). The characters employed in the
present analysis result in a sequence of relationships
between these taxa which differs from the solution
provided by Dufour. Compared with Dufour’s clado-
gram, corsosignata and haennii have changed places in
the present phylogeny, necessitating the elevation in
rank of haennii from subspecies of subsignata to spe-
cies (see Appendix A). Dufour substantiated his con-
cept of a monophyletic group containing corsosignata,
haennii, subsignata subsignata, and subsignata cazorla
by the degree of sclerotization and shape of the ven-
tromedial appendage of male sternite 8 and the de-
gree of sclerotization of female sternite 9 (Dufour
1991, characters 1 and 2). I can see no essential differ-
ences between the sclerotization and shape of the ven-
tromedial extension of male sternite 8 in this clade of
four taxa and that of other derived members of
Savtshenkia such as persignata, sardosignata, and sig-
nata, and prefer to consider it a plesiomorphy at this
level. The presence of an unsclerotized female sternite
9 as a synapomorphy for the members of the clade
haennii to subsignata cazorla has not been employed
in this paper because it is not always clear whether
sternite 9 should be valued as sclerotized or not in
species of Savtshenkia. Moreover, the female of sub-
signata cazorla remains unknown. Dufour considered
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
haennii the sister species of the pair subsignata subsig-
nata and subsignata cazorla because of the presence of
a ridge on what he denoted as the median part of the
inner gonostylus (Dufour 1991, character 3). As dis-
cussed under character 34, I consider the presence of
the anteriorly extended part of the lateral ridge of the
inner gonostylus an apomorphy of the pair subsignata
subsignata and subsignata cazorla. Of the three apo-
morphies of Dufour supporting the monophyly of
subsignata subsignata and subsignata cazorla, two ap-
pear to be invalid (Dufour 1991, characters 4 and 5).
The continuity of the posterior and ventral parts of
the lateral ridge of the inner gonostylus in these two
subspecies (Dufour 1991, character 4) probably is a
plesiomorphy (see discussion under character 34).
The relatively broad base of the outer gonostylus in
subsignata subsignata and subsignata cazorla (Dufour
1991, character 5) probably also represents a plesio-
morphy considering the situation in most related spe-
cies. The width of the base of the outer gonostylus
furthermore varies within the species and subspecies
and differs only slightly from that in /aennii.
Dufour’s third argument for the monophyly of the
pair subsignata subsignata and subsignata cazorla
(Dufour 1991, character 6, shape of the gonapoph-
ysis) is employed in this paper as well (character 14).
Following on his description of eleonorae,
Theischinger (1978) suggested that this species is
most closely related to benesignata. I am, however,
convinced that the sister species of eleonorae is boreo-
signata, which is substantiated by the almost identical
shape of the inner gonostylus (character 42) and the
bizarre modification of the aedeagal complex (charac-
ter 52). Both boreosignata and eleonorae are known of
their holotypes only, with the type localities lying
about 2500 kilometers apart, that of boreosignata in
northern Sweden and that of eleonorae in western
Rumania!
The clade benesignata to omega contains the species
which Theowald (1973) included in his signata
group, with the addition of the species omega and tri-
nacria described herein as new.
DISTRIBUTION
In this section I will give a brief account on the dis-
tribution of the species and subspecies of Savtshenkia.
As noted above, more detailed information on the
distribution of the western Mediterranean species and
subspecies and their closest relatives will be given in a
forthcoming paper on the historical biogeography of
the western Mediterranean area. A survey of the dis-
tribution of the species and subspecies of Savtshenkia
is given for the Afrotropical Region by Hutson (1980,
under Pterelachisus), the Nearctic Region by Alex-
ander (1965b), the Oriental Region by Alexander &
307
we
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Alexander (1973), and the Palaearctic Region by
Oosterbroek & Theowald (1992). The sequence of
taxa in the summary below follows that of fig. 2.
The two species of the pair alpha and grisescens dif-
fer considerably in their ranges. The first species is
known of its type locality in the Rodna Mountains of
northern Rumania only, while grisescens is distributed
over the central belt of Europe eastwards to the Altai
Mountains in central Asia.
The species of the clade hartigiana to simulans are
confined to the western Palaearctic, with most of
them having a restricted range. The species hartigiana
is known from Corsica and Sardinia only, atlas is re-
corded from Algeria and Morocco, and breviantenna-
ta from Spain, southern Switzerland, Corsica, Italy,
including Sardinia, and northern Algeria. The species
tulipa was described from two localities in southern
Switzerland, villeneuvii is known from central Spain,
while goriziensis, cheethami and simulans are more
widely distributed over the mountainous areas of cen-
tral Europe. The species ignobilis has an eastern
Nearctic distribution between about 35° and 50°N
latitude. The species subnodicornis has a wide range
extending from the Faeroes and Ireland in the west
over central Europe towards Lake Baykal in the east.
The clade baltistanica to chrysocephala contains spe-
cies which cover a wide geographic area. The species
baltistanica occurs in the Caucasus, Tadzhikistan and
Kashmir, glaucocinerea has an arctic Siberian distribu-
tion, as does its presumed sister species postposita,
which is furthermore known from northern Finland,
the Altai Mountains and Mongolia. The species
lundbladi is an endemic of Madeira, the species mul
tipicta is presently recorded from the Canary Islands
only. The nominotypical form of rufina is wide-
spread, ranging from Iceland and north-west Africa
towards Tadzhikistan in the east, while the subspecies
rufina maderensis is endemic to Madeira. The major-
ity of species of the grouping nephrotomoides to chry-
socephala is distributed in the mountainous areas sur-
rounding Lake Victoria in eastern Africa. The only
member of this group that does not occur in this re-
gion is the species draconis from South Africa (Natal).
The species alpium has a wide distribution which
includes most of Europe. It also occurs along the east-
coast of the Nearctic Region between about 42° and
55° N latitude (see Appendix A, synonymy, under
nebulipennis). The species macaronesica is endemic to
the Azores.
As is presently known, the phylogenetically rather
isolated species jeekeli has a disjunct distribution
throughout the Mediterranean, ranging from Galicia
in north-west Spain to Cyprus in the east.
The species obsoleta is distributed over most of
Europe, extending its range towards Georgia in the
south-east, while the pair holoptera and pagana is con-
308
fined to the central belt of Europe, with holoptera
having a disjunct distribution in south-west England,
north-west Spain, and Czechia.
Of the species of the clade confusa to aster, confusa
occurs over most of Europe, including the Faeroes
and Ireland in the north-west. The species interserta
has a distribution ranging from the central European
mountains in the west towards the Tuva in the east.
The beautiful ornata was described on the basis of a
single male from northern Israel, but a series of this
species, including females, was collected recently on
Cyprus by Pjotr Oosterbroek and Cita Hartveld. It
seems that the species graciae still is known from its
type locality in southern California only. Both sordi-
dipes and venerabilis are restricted to Kashmir. The
species eugeni is presently known from Corsica, Sicily,
and southern Italy. The species serrulifera has a more
extensive distribution, occurring in mountainous are-
as from the Pyrenees in the west to the Caucasus in
the east. Theowald (1973) described aspromontensis
from southern Italy (Aspromonte), but more recently
it was collected also in Greece (Peloponnisos, Crete).
The species aster still is known of its holotype only,
originating from the Greek island Thasos in the
northern Aegean Sea.
The presumed sister species fragilis and phoroctenia
have a Nearctic distribution which extends across the
continent in the region of the 50th degree of latitude.
The species odontostyla is known from the north-east-
ern coastal areas of the Black Sea and the Greek island
Thasos.
The species fragilina has a Nearctic distribution,
being recorded from the north-western part of the
continent, ranging from Alaska to Colorado, as well
as from Maine on the east coast (under its synonym
broweri). The species limbata is widespread in the
mountainous areas of Eurasia, subvafra seems to be
confined to continental western Europe. The mem-
bers of the clade pechlaneri to gimmerthali pteromacu-
lata are distributed in the mountainous areas of the
western Palaearctic Region, with pechlaneri having a
disjunct distribution from Andorra in the Pyrenees to
northern Iran, gimmerthali gimmerthali ranging from
the Sierra Nevada of Spain in the south towards
northern Scandinavia and eastwards to the Caucasus,
gimmerthali mattheyi being restricted to southern
France, and gimmerthali pteromaculata to Corsica.
The three subspecies of invenusta differ consider-
ably in the extent of their distribution area. The sub-
species invenusta microinvenusta is presently known
only from southern Spain (Sierra Nevada), the sub-
species invenusta subinvenusta has a wider range in the
mountains of central Europe, while the nominotypi-
cal subspecies disjunctly ranges across Eurasia and al-
so occurs in eastern North America (see Appendix A,
synonymy, under insignifica and packardi).
The species cyrnosardensis is endemic to Corsica
and Sardinia, staegeri occurs in Europe from the
Faeroes to northern Spain in the west towards the
coastal areas of the Baltic Sea in the east. Both species
of the pair nielseni and sardosignata have a restricted
range, with nielseni being confined to the Alps, and
sardosignata occurring on Corsica and Sardinia, as
well as in southern France.
The four members of the clade haennii to subsigna-
ta cazorla are also restricted to Europe, with haennii
being recorded from the French and Spanish
Pyrenees, corsosignata occurring on Corsica, subsigna-
ta cazorla in northern Spain, and subsignata subsigna-
ta being more widely distributed in the mountains of
central Europe. As mentioned above, both boreosig-
nata and eleonorae are known of their holotypes only,
with the type localities lying in northern Sweden and
western Rumania respectively. The distribution area
of the species benesignata consists of widely separated
populations ranging from the Alps towards northern
Scandinavia and eastwards to the Tien Mountains.
The original descriptions of the two subspecies persig-
nata persignata and persignata tofina were based on
material originating from North Korea, to which area
the distribution range of the nominotypical form
seems to be restricted. Later records of persignata toft-
na, however, showed that this subspecies ranges
across all of northern Eurasia.
The species signata is widespread in Europe, cover-
ing most of the area. The triplet trinacria to omega is
restricted to Italy, with ¢rimacria occurring on Sicily,
mannheimsi in the north of Italy, and omega on
Sardinia.
ACKNOWLEDGEMENTS
For the loan of material of Savtshenkia I am most
indebted to the following persons: Roy Danielsson
(Lund), Christophe Dufour (Neuchatel), Jon Gelhaus
(Philadelphia), Brian Pitkin (London), Hans-Peter
Tschorsnig (Stuttgart), and Holly Williams (Was-
hington D.C.). Lali Eiroa (Santiago de Compostela,
Spain) kindly donated material of Savtshenkia holopte-
ra to the collection of the Zoological Museum,
Amsterdam. I would like to thank Wayne Mathis and
Holly Williams for their much appreciated help du-
ring my short visit to the United States National
Museum of Natural History, Washington D.C.
A draft version of this paper was read and com-
mented upon by Pjotr Oosterbroek, Hans Duffels,
Fred Schram, and an anonymous reviewer. Their crit-
icism has led to a number of substantial improve-
ments of the text. The investigations were supported
by the Life Science Foundation (SLW), which is sub-
sidized by the Netherlands Organization for
Scientific Research (NWO).
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
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on genital characters. -Beaufortia 35: 135-174.
Theischinger, G., 1978. Tipula (Savtshenkia) eleonorae
sp.nov. aus Rumänien (Diptera, Tipulidae). —
Entomologische Berichten 38: 45-46.
Theischinger, G., 1983. Uber eine Aufsammlung von
Savtshenkia Alexander, 1965, aus dem östlichen
Mittelmeerraum (Diptera: Tipulidae). — Entomologische
Berichten 43: 26-28.
Theowald, B. 1957. Die Entwicklungsstadien der Tipuliden
(Diptera, Nematocera), insbesondere der west-palaeark-
tischen Arten. — Tijdschrift voor Entomologie 100: 195-
308.
Theowald, B. 1967. Familie Tipulidae. (Diptera, Nema-
tocera). Larven und Puppen. — Bestimmungsbiicher zur
Bodenfauna Europas 7: 1-100.
Theowald, B., 1973. Tipulidae. — Fliegen der Palaeark-
tischen Region 15, Lieferung 300: 321-404.
Theowald, B., 1978. Tipulidae. — Fliegen der Palaeark-
tischen Region 15, Lieferung 318: 405-436.
Theowald, B. & C. Dufour, 1983. Tipula (Savtshenkia)
gimmerthali mattheyi ssp.n. from the French Pyrenees. —
Mitteilungen der Schweizerischen Entomologischen
Gesellschaft 56: 283-284.
Theowald, B., C. Dufour & P. Oosterbroek, 1982. The
zoogeography of the western Palaearctic Tipulidae
(Diptera). Part IV: the Tipulidae of Corsica and Sardinia
with a note on Dolichopeza fuscipes Bergroth. —
Mitteilungen der Schweizerischen Entomologischen
Gesellschaft 55: 317-332.
Theowald, B. & P. Oosterbroek, 1981. Zur Zoogeographie
der westpalaearktischen Tipuliden II. Die Tipuliden der
iberischen Halbinsel (Diptera, Tipulidae). — Beaufortia
30: 179-192.
Wood, H. G., 1952. The crane-flies of the South-West
Cape (Diptera, Tipuloidea). — Annals of the South
African Museum 39: 1-327.
Received: 2 June 1994
Accepted: 13 September 1994
APPENDIX A
In this appendix taxonomic changes pertaining to
Savtshenkia are dealt with. The changes include the
description of three new western Palaearctic species of
the subgenus, the establishment of five synonymies,
the elevation of one subspecies to species rank, and
the removal of one species from Savtshenkia.
Descriptions of new species
Tipula (Savtshenkia) alpha sp. n.
(figs. 152-170)
Type material. — Holotype d: Rumania, Mara-
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 152-164, Tipula (Savtshenkia) alpha sp. n. — 152, male head and appendages, lateral view; 153, male head, dorsal view,
antennae and mouthparts omitted; 154, female antenna, basal segments; 155, male terminalia, lateral view; 156, male tergi-
te 9-10, dorsal view; 157, male tergite 9-10, posterior view; 158, male sternite 8, posterior margin, ventral view; 159, right
sp2, dorsal view; 160, left outer gonostylus, lateral view; 161, left inner gonostylus, lateral view; 162, aedeagal guide with right
gonapophysis and fragmentum, posteromedial view; 163, sperm pump and appendages, lateral view; 164, sperm pump and
appendages, anterior view.
mures, Muntii Rodnei, Borsa, subalpine lake
‘Stiolul’, main sources of river Bistrita Aurie, 1700-
1800 m, 31.V.1974, L. Botosaneanu (ZMAN). —
Paratypes 5d, 19, same data as holotype (ZMAN).
Material preserved in alcohol.
Description
Body length 10-12 mm (dg), 14 mm (2); wing
length 12-13.5 mm (dé), 13.5 mm (@); antennal
length 5-5.5 mm (4), ? mm (9 ; antenna broken, fig.
154).
312
Colour. — Ground colour greyish brown. Head and
thorax dark greyish. Antenna with scapus dark grey-
ish, remainder of segments dark brown. Palpus dark
brown. Prescutum with single broad medial and pair
of lateral dark stripes. Wing veins brown; pterostigma
brown, distinct; membrane unpatterned. Haltere yel-
lowish brown, including knob. Trochanters of same
dark greyish colour as coxae, femora yellowish brown
in basal section, from about half length darkening to-
wards tip, remainder of legs dark brown. Abdomen
dark brown, terminalia slightly darker. (Colour de-
165
ET NS
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
Figs. 165-170. Tipula (Savtshenkia) alpha sp. n., female. — 165, ovipositor, lateral view; 166, sternite 8 and hypogynial valves,
ventral view; 167, sternite 8 and hypogynial valves, dorsal view; 168, sternite 9, medial part, ventral view; 169, genital fork,
dorsal view; 170, spermathecae.
duced by approximation, material preserved in alco-
hol).
Head (figs. 152-154). — Rostrum shorter than re-
mainder of head, nasus well developed; eyes small,
dorsally and ventrally separated by about four times
diameter of scapus; occiput perpendicular to lateral
sides of head (dorsal aspect, fig. 153); male antenna
about four times as long as head, longest verticils
about one and a half times as long as diameter of fla-
gellomeres at base; female antenna shorter (flagella of
both antennae broken beyond basal flagellomeres in
female paratype, fig. 154).
Thorax. — Wing fully developed in both sexes,
squama devoid of macrotrichia; cell ml gradually
widening towards wing margin. Tarsal claws tooth-
less in both sexes.
Male terminalia (figs. 155-164). — Compactly built,
tergite 9-10 and gonocoxites separated by membra-
nous zone (fig. 155). Tergite 9-10 (figs. 156, 157)
broad and short, with medial membranous area occu-
pying about one third of total width; posterior margin
widely V-shaped, with shallow emargination near me-
dian line; margin ventrally set with scattered hairs,
without spines; lateral corners broadly rounded.
Posterior margin of sternite 8 unmodified (fig. 158).
Gonocoxites ventrally separated by membranous area,
membrane with pair of weakly sclerotized bands ex-
tending anteriorly from base of aedeagal guide. Suture
x narrow, about one fourth length of gonocoxite (fig.
155). Sp2 (fig. 159) long, medial blackish sclerotized
margin moderately serrate; dorsally separate from ter-
gite 9-10; connexion between sp2 and posterior mar-
gin of foramen of gonostyli short (fig. 159, arrow).
Outer gonostylus (fig. 160) broad, its tip anteriorly in-
clined, posterior margin only moderately blackened.
Inner gonostylus (fig. 161) with relatively slender
blackish sclerotized anterior part; lateral rim in its ven-
tral part directed towards base of gonostylus; postero-
laterally near base with blackened rim (fig. 161, ar-
row). Aedeagal guide (fig. 162) a short and plump
blackish sclerotized structure, gonapophysis reduced
to a sclerotized strip (fig. 162, arrow). Fragmentum
(fig. 162) a rounded lobe with a few long hairs dorso-
medially. Sperm pump (figs. 163, 164) with anterior
apodemes short, posterior apodemes well developed
and obliquely placed, compressor apodeme dorsally
emarginate, blackish; lumen of endophallus well de-
veloped; ventral wall convex. Aedeagus short, anterior-
ly reaching segment 8 (fig. 155), tubular throughout,
gradually narrowing towards tip.
Female terminalia (figs. 165-170). — Cercus and
hypogynial valve well developed, robust, relatively
short (fig. 165). Sternite 8 dorsolaterally near base of
hypogynial valve with concavity (fig. 165, arrow);
ventral membrane in posterior part of sternite 8 en-
closing medial base of hypogynial valves (fig. 166).
Dorsomedial margin of hypogynial valve only moder-
ately blackened (fig. 167). Sternite 9 with slender me-
dial projection (fig. 168). Genital fork elongate, ante-
riorly broader (fig. 169). Three spermathecae,
globular to oblong, of about same size, moderately
sclerotized (fig. 170).
Etymology. — The name of this species was
prompted by its phylogenetic position within
Savtshenkia and is the pendant of Tipula
(Savtshenkia) omega sp. n. A noun in apposition.
Remarks. — Tipula (Savtshenkia) alpha differs from
its presumed sister species 7. (S.) grisescens by the
315
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
x 173
176
Figs. 171-176. Tipula (Savtshenkia) omega sp. n., male. — 171, head and appendages, lateral view; 172, head, dorsal view, an-
tennae and mouthparts omitted; 173, terminalia, lateral view; 174, tergite 9-10, dorsal view; 175, tergite 9-10, posterior view;
176, left sclerotization proctiger, lateral view.
colour of the coxae (dark greyish in alpha, yellowish
brown in grisescens), the less elongate male antenna
(see figs. 9 and 152), the separated gonocoxites (fused
in grisescens), the broader outer gonostylus (see figs.
76 and 160), the more slender anterior part of the in-
ner gonostylus and the presence of a sclerotized rim
posterolaterally at the base of the same structure, the
shallowly emarginate compressor apodeme of the
sperm pump (deeply emarginate in grisescens), the
rounded body of the sperm pump (with conical ante-
roventral extension in grisescens).
The type locality of alpha lies above the tree line on
the subalpine plain at an altitude of 1700-1800 m.
314
The locality has a characteristic vegetation of Pinus
mugo Turra and the grass Nardus stricta Linnaeus. At
the end of May thaw has set in, but the surface of the
area is still extensively snow- and ice-covered. Under
sunny conditions alpha emerges in large numbers as
soon as holes begin to appear in the ice-covering.
Although fully winged, the insects tend to crawl
around on the ice and surrounding grasses. Under
cloudy conditions alpha shows no sign of activity,
suggesting that the behaviour of the species is largely
influenced by the environmental temperature. In
these aspects, the biology of alpha resembles that of
the trichopteron Chionophylax czarnohoricus Dziedz.,
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
Figs. 177-184. Tipula (Savtshenkia) omega sp. n., male. — 177, sternite 8, posterior margin, ventral view; 178, sternite 8, pos-
terior margin, posterior view; 179, right sp2, dorsal view; 180, left outer gonostylus, lateral view; 181, left inner gonostylus,
lateral view; 182, aedeagal guide with right gonapophysis and fragmentum, posteromedial view; 183, sperm pump and ap-
pendages, lateral view; 184, sperm pump and appendages, anterior view.
a species which occurs in the same locality (Dr
Botosaneanu, pers. comm.).
Tipula (Savtshenkia) omega sp. n.
(figs. 171-184)
Type material. — Holotype d: Italy, Sardinia,
Fiume Taloro, 670 m, 9°31/40°06, 21.X.1981, H.
Malicky (ZMAN). — Paratypes: 1d, Italy, Sardinia, Rio
Aratu, 970 m, 9°15/40°02, 22.X.1981, H. Malicky
(ZMAN). Material preserved in alcohol.
Description
Body length 12-14.5 mm (d); wing length 14-
15.5 mm (d); antennal length 5-6 mm (d).
Colour. — Ground colour yellowish brown. Vertex
of head and dorsal and lateral sides of thorax darker,
probably slightly greyish pruinose in dry specimens.
Antenna yellowish brown, scapus and pedicel some-
what lighter coloured than flagellum. Palpus yellow-
ish brown. Presutum with confluent medial and pair
of lateral dark stripes indicated. Wing veins yellowish
brown; pterostigma yellowish brown, distinct; mem-
brane vaguely patterned. Halteres yellowish brown,
including knob. Legs yellowish brown, femora and
tibiae darkened at extreme tip, tarsi darkening to-
wards tip. Abdomen yellowish at base, darkening to-
wards tip. (Colour deduced by approximation, mate-
rial preserved in alcohol).
Head (figs. 171, 172). — Rostrum about as long as
remainder of head, nasus well developed; eyes well
developed, dorsally separated by about two and a half
times diameter of scapus, ventrally by about two
times diameter of scapus; occiput gradually narrow-
ing towards cervix (dorsal aspect, fig. 172); male an-
tenna just less than two times length of head, longest
verticils about as long as flagellomeres (fig. 171).
Thorax. — Wing fully developed, squama devoid of
macrotrichia; cell m1 elongate bell-shaped, narrow-
ing towards wing margin. Tarsal claws with small me-
dial teeth in male.
Male terminalia (figs. 173-176). — Large, tergite 9-
GIS
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 185-191. Tipula (Savtshenkia) trinacria sp. n., male. — 185, terminalia, lateral view; 186, tergite 9-10, dorsal view; 187,
tergite 9-10, posterior view; 188, left sclerotization proctiger, lateral view; 189, sternite 8, posterior margin, ventral view; 190,
sternite 8, posterior margin, posterior view; 191, right sp2, dorsal view.
10 and gonocoxites separated by membranous zone
(fig. 173). Tergite 9-10 (figs. 174, 175) elongate,
with narrow medial membranous area; posterior mar-
gin with pair of broad laterally diverging extensions;
extensions not blackish sclerotized, devoid of spines
along ventral margin. Proctiger with rather diffuse
and narrow lateral sclerotization (fig. 176). Sternite 8
(figs. 173, 177, 178) posteriorly with diverging broad
dorsoventrally flattened lobes, each lobe carrying
about 35-40 black spines. Gonocoxites ventrally
fused with narrow medial carina; at level of insertion
of inner and outer gonostyli with concentration of
316
long and strong setae (fig. 173). Suture x narrow,
about two-fifth length of gonocoxite (fig. 173). Sp2
(fig. 179) medially protruding, blackish sclerotized
medial margin entire; sp2 fused with tergite 9-10;
connexion between sp2 and posterior margin of fora-
men of gonostyli long. Outer gonostylus (fig. 180)
with elongate slender anterior part, posterior blackish
sclerotized part relatively broad, with short poste-
rodorsal extension, slightly concave. Inner gonostylus
(fig. 181) with anterior and posterior parts about as
long, posterior part caudally terminating in a densely
pubescent lobe. Aedeagal guide (fig. 182) long and
H. DE Jonc: Phylogeny of Tipula (Savtshenkia)
Figs. 192-196. Tipula (Savtshenkia) trinacria sp. n., male. — 192, left outer gonostylus, lateral view; 193, left inner gonosty-
lus, lateral view; 194, aedeagal guide with right gonapophysis and fragmentum, posteromedial view; 195, sperm pump and
appendages, lateral view; 196, sperm pump and appendages, anterior view.
slender, blackish sclerotized, gonapophysis short,
dorsally directed. Fragmentum (fig. 182) anteriorly
with sclerotized bar which dorsally carries a bundle of
long hairs, posteriorly weakly sclerotized. Sperm
pump (figs. 183, 184) with large anterior apodemes,
posterior apodemes caudally directed, compressor ap-
odeme dorsally deeply emarginate, wings broad; lu-
men endophallus small, body wall ventrally concave.
Aedeagus elongate (fig. 173), anteriorly reaching seg-
ment 3, tubular throughout.
Etymology. — The phylogenetic position of this
species within Savtshenkia suggested its name. A
noun in apposition.
Remarks. — The species omega very much resembles
the species mannheimsi and trinacria sp. n. It differs
from these species by the number of spines on the
posterior lobes of male sternite 8 (ca. 35-40 in omega
to ca. 50 in mannheimsi and ca. 65 in trinacria), the
absence of spines on the ventral margin of male ter-
gite 10 in omega (spines absent also in mannheimsi,
present in #rinacria), and in the shape of the inner and
outer gonostyli.
The female of omega remains unknown.
Tipula (Savtshenkia) trinacria sp. n.
(figs. 185-202)
Type material. — Holotype d: Italy, Sicily,
Nébrodi, 10 km north of Capizzi, Fiume Troina,
1200-1400 m, 11-14.X.1993, P. Oosterbroek & C.
Hartveld (ZMAN). — Paratypes: 135,32, same data as
holotype; 46 , Italy, Sicily, Nébrodi, 5 km north of
Monte Soro, Biviere di Cesaro, 1300 m, 12.X.1993,
P. Oosterbroek & C. Hartveld; 2d, Italy, Sicily,
Nébrodi, 6 km east of Serra del Re, Foresta Vécchia,
Fiume di Saracena, 1100 m, 13.X.1993, P.
Oosterbroek & C. Hartveld; 4d, 39, Italy, Sicily,
Madonie, 6 km north of Monte San Salvatore,
Torrente Vicaretto, 1300 m, 15-21.X.1993, P.
Oosterbroek & C. Hartveld, (ZMAN).
Description
Body length 13.5-16 mm (d), 15.5-17 mm (2);
wing length 16-19 mm (dé), 16.5-18 mm (9); anten-
nal length 6-7 mm (d), 5-5.5 mm (9).
Colour. — Ground colour yellowish brown.
Excepting rostrum, head greyish pruinose. Antenna
with yellowish scapus and pedicel, flagellum almost
uniformly brownish, basal flagellomeres somewhat
lighter. Palpus yellowish brown. Dorsal and lateral
sides of thorax slightly greyish pruinose. Presutum
DIV
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Figs. 197-202. Tipula (Savtshenkia) trinacria sp. n., female. — 197, ovipositor, lateral view; 198, sternite 8 and hypogynial val-
ves, ventral view; 199, sternite 8 and hypogynial valves, dorsal view; 200, sternite 9, medial part, ventral view; 201, genital
fork, dorsal view; 202, spermathecae.
with pair of approximating slender medial and pair of
lateral dark stripes, medial stripes anteriorly lighter
coloured. Wing veins brown; pterostigma brown, dis-
tinct; membrane vaguely patterned and iridescent.
Halteres with yellowish brown pedicel, knob brown.
Legs yellowish brown, femora and tibiae darkened at
extreme tip, tarsi largely dark brown. Abdomen yel-
lowish at base, darkening towards tip.
Head. — Very much as that of omega sp. n. (see figs.
171, 172; see for description of head under that spe-
cies, above).
Thorax. — Wing fully developed in both sexes,
squama devoid of macrotrichia; cell m1 elongate bell-
shaped, narrowing towards wing margin. Tarsal claws
with small medial teeth in the male, female claws
toothless.
Male terminalia (figs. 185-196). — Large, tergite 9-
10 and gonocoxites separated by membranous zone
(fig. 185). Tergite 9-10 (figs. 186, 187) elongate,
318
with narrow medial membranous area; posterior mar-
gin with pair of broad laterally diverging extensions;
extensions blackish sclerotized along ventral margin,
ventral margin set with small black spines. Proctiger
with rather well demarcated slender lateral sclerotiza-
tion (fig. 188). Sternite 8 (figs. 189, 190) posteriorly
with diverging broad flattened lobes, each lobe carry-
ing about 65 slender black spines. Gonocoxites ven-
trally fused with narrow medial carina. Suture x nar-
row (fig. 185). Sp2 (fig. 191) medially protruding,
blackish sclerotized medial margin entire; sp2 fused
with tergite 9-10; connexion between sp2 and poste-
rior margin of foramen of gonostyli long. Outer go-
nostylus (fig. 192) with elongate slender anterior part,
posterior blackish sclerotized part elongate, upcurved.
Inner gonostylus (fig. 193) with anterior and posteri-
or parts about as long, posterior part anteriorly ex-
tended, posteriorly terminating in cup-shaped struc-
ture. Aedeagal guide (fig. 194) long and slender,
blackish sclerotized, gonapophysis short, dorsally di-
rected. Fragmentum (fig. 194) anteriorly with sclero-
tized bar which dorsally carries a bundle of long hairs,
posteriorly weakly sclerotized. Sperm pump (figs.
195, 196) with large anterior apodemes, posterior ap-
odemes caudally directed, at tip somewhat upturned,
compressor apodeme dorsally deeply emarginate,
with narrow cleft, wings broad; lumen endophallus
small, wall ventrally concave. Aedeagus elongate (fig.
185), anteriorly reaching segment 4, tubular through-
out.
Female terminalia (figs. 197-202). — Cercus and
hypogynial valve well developed, cercus approximate-
ly straight (fig. 197). Ventral membrane in posterior
part of sternite 8 terminating in between bases of hy-
pogynial valves (fig. 198). Dorsomedial margin of hy-
pogynial valve only moderately blackened (fig. 199),
ventrally at base with concavity (figs. 197, 198).
Sternite 9 with relatively narrow and slender medial
projection, concave lateral of this structure (fig. 200).
Genital fork short and broad, broadest at anterior side
(fig. 201). Three spermathecae, globular, of about
same size, blackish sclerotized (fig. 202).
Etymology. — Trinacria is an old name of Greek or-
igin for Sicily. A noun in apposition.
Remarks. — The species trinacria very much resem-
bles mannheimsi and omega sp. n. For distinguishing
characters see under omega. The type specimens of
trinacria were captured along rivers and a lake in de-
ciduous forests. The specimens collected in the peri-
od from 15-21 October were taken from the moss
covered river banks of the Torrente Vicaretto. Among
these latter specimens was one pair in copula.
Synonymy
insignifica Alexander, 1924 = invenusta invenusta
Riedel, 1919 syn. n.
Alexander (1924) described insignifica as a new
species based on a single male originating from
Mount Washington, New Hampshire, U.S.A.
Alexander (loc. cit.) recorded the collection of BSNH
as the depository of the holotype (holotype not exam-
ined). I have studied specimens labelled by Alexander
as metatypes, i.e. specimens compared with the type
specimen(s), originating from Mount Katahdin,
Maine and Mount Madison, New Hampshire (-
USNM). Examination of these specimens revealed only
slight differences between the Nearctic insignifica and
the Palaearctic invenusta invenusta in the distance of
the eyes on the ventral side of the head (see also
‘Discussion of adopted phylogeny’). No other struc-
tural differences between insignifica and invenusta in-
venusta were found and I therefore consider insignifi-
H. DE Jone: Phylogeny of Tipula (Savtshenkia)
ca a junior synonym of invenusta invenusta. [Contrary
to Alexander’s (1942, 1966b) assertion, the posterior
margin of tergite 9-10 is posteroventrally armed with
small black spines in insignifica, as it is in invenusta].
broweri Alexander, 1940 = fragilina Alexander, 1919
syn. n.
Alexander (1940) described broweri based on 12
males and two females from Mount Katahdin,
Maine, U.S.A. (types in usNM). Despite Alexander’s
claim that broweri should be entirely distinct from its
relatives, examination of type material of broweri
showed that it is morphologically identical with frag
ilina. | therefore consider broweri a junior synonym
of fragilina.
nebulipennis Alexander, 1919 = alpium Bergroth,
1888 syn. n.
Alexander (1919a) introduced nebulipennis as a new
species based on two males from Battle Harbour,
Labrador, Canada (holotype in Brooklyn Museum,
New York, U.S.A., paratype in USNM). Later, nebuli-
pennis was also recorded from Quebec and New
Hampshire (Alexander 1965b). Examination of mate-
rial of nebulipennis in the Alexander collection (USNM)
revealed no significant morphological differences
between these specimens and the European species at
pium. | therefore consider nebulipennis a junior syno-
nym of alpium. The larvae of alpium were recorded
from terrestrial mosses (Brindle 1958, 1959, 1960).
Theowald (1957, 1967) recorded the larvae further-
more from the litter layer beneath oaks. Possibly a/pi-
um has been introduced into North America in a way
similar to the European species Tipula (Tipula) palu-
dosa Meigen (see Alexander 1965a).
packardi Alexander, 1928 = invenusta invenusta
Riedel, 1919 syn. n.
Alexander (1928) described Tipula packardi based
on a single male from Hopedale, Labrador, Canada.
The holotype of packardi is preserved in cncı (holo-
type not examined). An additional male specimen
originating from the type locality of packardi is
present in USNM. This latter specimen was compared
by Alexander with the holotype of packardi and con-
sidered conspecific. Examination of the metatype of
packardi in USNM showed that it has the eyes on the
ventral side of the head slightly more approximate
than the examined specimens of invenusta invenusta
(see also ‘Discussion of adopted phylogeny’, and ##-
significa, this section). No other structural differences
were found and I therefore consider packardi a junior
synonym of invenusta invenusta.
perparvula Alexander, 1926 = ignobilis Loew, 1863
syn.n. (partim?)
319
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Loew (1863) described the species ignobilis based
on the female sex from material captured by Osten-
Sacken in the District of Columbia, U.S.A. (holotype
in MCZC, not examined). According to its labels,
Alexander compared a male specimen captured by
Osten Sacken in Catskill, New York, with the type
specimen of ignobilis and considered the two conspe-
cific. Examination of this inferred male specimen of
ignobilis and the paratype male of the species perpar-
vula (both in USNM) revealed that these two speci-
mens belong to the same species. Alexander (1926),
describing perparvula based on two males originating
from Aweme, Manitoba, Canada (holotype in CNCI,
paratype in USNM), explicitly stated that perparvula
has a relatively stout nasus. Examination of the para-
type of perparvula shows that this structure is entirely
absent in the specimen, as it is in the species zgnobilis.
Either the holotype and paratype of perparvula be-
long to two different species, with the paratype being
conspecific with zgnobilis, or both type specimens are
conspecific and belong to ignobilis. Examination of
the holotype of perparvula should settle this question.
Subspecies raised to species rank
haennii Dufour, 1991
This taxon was originally described as a subspecies
of subsignata (Dufour 1991). The tip of the aedeagal
guide of haennii does not show the abrupt constric-
tion which probably represents a synapomorphy of a
monophyletic group containing corsosignata, subsig-
nata subsignata and subsignata cazorla (character 8,
above). Because of its phylogenetic position next to
this clade, haennii is elevated from subspecific to spe-
cific rank.
Species removed from Savtshenkia
convexifrons Holmgren, 1883
Savchenko (1961) placed the species convexifrons
Holmgren in his subnodicornis group of Prerelachisus
sensu Savchenko, nec Rondani. Savchenko (1961:
223) based its membership of the subnodicornis group
on the shape of the inner gonostylus, and its incorpo-
ration in Pterelachisus sensu Savchenko on the proxi-
mal position of the fork of vein M3+4. At the same ti-
me, Savchenko underlined the isolated position of
convexifrons within this grouping. As noticed in the
introduction to the present paper, the name
Savtshenkia replaces Pterelachisus sensu Savchenko.
Theowald (1973) followed Savchenko in considering
convexifrons a species of Savtshenkia, but split the sub-
nodicornis group sensu Savchenko into two species
groups (table 3). I remove convexifrons from
Savtshenkia because of the absence of the defining ap-
omorphies 53 and 83 of Savtshenkia. Furthermore,
320
the species convexifrons differs from the species of
Savtshenkia by the shape of male tergite 9-10, the
ventral structure of the fused gonocoxites, the shape
of the elongate aedeagal guide and its gonopophyses,
and, most noteworthy, by the presence of a complete
genital bridge (a plesiomorphy; the medial part of the
genital bridge is absent in all species of Savtshenkia).
The fork of vein M3+4 is located proximally in many
genera and subgenera of Tipulidae other than
Savtshenkia. The resemblance of the inner gonostylus
of convexifrons to the inner gonostyli of plesiomorp-
hous species of Savtshenkia is only superficial. At the
moment I am unable to establish the actual phyloge-
netic relationships of convexifrons and can not allocate
it to any other subgenus or species group of Tipulidae.
APPENDIX B
In this appendix are listed the species and subspe-
cies of Savtshenkia which, because of deficiency of da-
ta, were not incorporated in the character state matrix
given in table 4. Incompleteness of data resulted of
several causes. For some of the pertaining taxa no ma-
terial was examined, for others the material examined
was incomplete as regards the structures of the male
terminalia, for others again only the female is known,
thus excluding their incorporation in a phylogeny
which is largely based on characters of the male ter-
minalia. Where possible, the presumed phylogenetic
position of the species and subspecies concerned is in-
dicated. For the deposition of the type material of
species described by Savchenko see Kandybina,
Lantzov & Savchenko (1987).
Unexamined
imperfecta Riedel, 1914. — Description based on
single brachypterous male specimen from Kenya
(Mount Kenya) and single brachypterous female spe-
cimen, possibly belonging to the same species, from
Tanzania (Mount Kilimanjaro). Types in MNHN.
Species doubtfully belongs to Savtshenkia. Distribu-
tion: Kenya, Tanzania.
letifera Alexander, 1951. — Description based on
single male from China (Tibet, Shugden Gompa a.k.a.
Hsiu-teng). Type in BMNH. Inner and outer gonostyli
quite similar to those of Kashmirian sordidipes and
venerabilis. Differences with these species in structures
of posterior margins of tergite 9-10 and sternite 8.
Shape of sternite 8 suggests relationship with Nearctic
graciae and Palaearctic kiushiuensis and koreana.
Alexander (1953) illustrated details male terminalia of
letifera. Distribution: China (south-east Tibet).
minuscula Savchenko, 1971. — Originally described
as Tipula (Savtshenkia) nana by Savchenko (1968a)
based on two males and single female from USSR
(Kamchatka). Types in zıas. Savchenko, probably
correctly, considered minuscula closely related to
Nearctic ignobilis, with which it should differ in
colour characteristics. Details male terminalia of mz-
nuscula illustrated by Savchenko (1968a).
Distribution: Russia (Kamchatka).
nivalis Savchenko, 1961. — Description based on
three males and four females from USSR
(Stavropolskiy kray and Gruzinskaya SSR). Types in
zias. Savchenko (1961) compared nivalis with glau-
cocinerea. Theowald (1973) considered both very
closely related and suggested that nivalis might be
considered a subspecies of glaucocinerea. Details male
terminalia of nivalis illustrated by Savchenko (1961)
and Theowald (1973). Distribution: Russia
(Stavropolskiy kray), Georgia.
productella Alexander, 1928. — Description based
on single male from Canada (Labrador). Type in
encı. According to Alexander (1928) most closely al-
lied to Nearctic fragilina. Structure of sternite 8 of
productella of same build as found in clade fragilina to
subvafra of present paper; productella probably be-
longs to this clade; synonymy with one of its species
possible. Distribution: Canada (Labrador).
subalpium Savchenko, 1961. — Description based
on single male from USSR (Krasnodarskiy kray,
Glavniy Mountains). Type in z1as. Savchenko (1961)
compared subalpium with alpium, with which it
shares same general colour and structure of inner and
outer gonostyli and to which it is probably closely re-
lated. Differs quite remarkably from alpium in struc-
ture of posterior margin of sternite 8: whereas poste-
rior margin of sternite 8 is U-shaped emarginate and
carries spines along its edge in alpium (fig. 105),
emargination is filled up with membrane covered
with short setae in subalpium. According to
Savchenko, there is a short pubescent extension in
between what he denotes sternites 8 and 9. This
structure could well be homologous with midventral
extension found in alpium and macaronesica, which
in dry material is often hardly visible or seems to orig-
inate from posterior margin of sternite 8 (see fig. 39).
Savchenko (1969) depicted male terminalia of sub-
apium in lateral and posterior view. Distribution:
Russia (Krasnodarskiy kray), Georgia.
tetragramma Edwards, 1928. — Description based
on single male from China (Yunnan). Type in BMNH.
Edwards (1928) was of the opinion that tetragramma
must bear a rather strong resemblance to kiushiuensis.
Distribution: China (Yunnan).
Incomplete male terminalia
kiushiuensis Alexander, 1925. — Description based
on single male and single female from Japan (Kyushu,
Mount Kirishima). Types in usnm. Glass slides with
terminalia male holotype and additional male speci-
men (USNM) do not allow study of all relevant struc-
H. DE JONG: Phylogeny of Tipula (Savtshenkia)
tures. Related to species in range jeekeli to aster of cla-
dogram fig. 2. Posteriorly produced unarmed medial
part of posterior margin male sternite 8 suggests rela-
tionship with koreana. Savchenko (1968b) illustrated
details male terminalia of kiushiuensis. Distribution:
Russia (Primorskiy kray), Japan (Kyushu, Shikoku).
koreana Alexander, 1934. — Description based on
single male and four females from North Korea
(Mount Kongo). Types in usnm. Glass slide with ter-
minalia male holotype does not allow study of all rel-
evant structures. Related to species in range jeekeli to
aster of cladogram fig. 2. Posteriorly produced un-
armed medial part of posterior margin male sternite 8
suggests relationship with kiushiuensis. Alexander
(1934) illustrated details male terminalia and wing of
koreana. Distribution: Russia (Sakhalin), North
Korea, Japan (Shikoku).
mohriana Alexander, 1954. — Description based on
single male and single female from Japan (Shikoku,
Mount Ishizuchi). Types in usnm. Glass slide with
terminalia male holotype does not allow study of all
relevant structures. Most probably sister species of
clade fragilina to subvafra of present paper as implied
by fused sp2 and tergite 9-10 (character 54), slender
anterior part outer gonostylus (character 45), bifid tip
of anterior part of inner gonostylus (character 30),
and widely emarginate bilobed and unarmed posteri-
or margin of male sternite 8 (cf. character 60).
Differing most notably from species of clade fragilina
to subvafra by presence of spinous gonapophysis (au-
tapomorphy) and absence of blackened dorsal margin
of fragmentum (cf. character 17). Details male termi-
nalia and wing of mohriana illustrated by Alexander
(1955). Distribution: Japan (Shikoku).
sciadoptera Alexander, 1964. — Described as guttu-
lifera by Alexander (1961) based on two males from
India (Himalayas). Types in usnm. Glass slides with
terminalia male holotype and paratype do not allow
study of all relevant structures. Probably related to
species in range baltistanica to chrysocephala of clado-
gram in fig. 2. Alexander (1961) illustrated details
male terminalia of sciadoptera (as guttulifera).
Distribution: India (Sikkim).
Known in female sex only
aberdareica ulugurica Alexander, 1962. -
Description based on single female from Tanzania
(Uluguru Mountains). Type in MRAC, not examined.
According to Alexander (1962) closely related to no-
minotypical form, sharing several colour characteris-
tics with chrysocephala. Distribution: Tanzania.
nephrotomoides invariegata Alexander, 1956. —
Description based on single female holotype from
Uganda (Ruwenzori Range). Type in BMNH.
Agreeing in its dark colour with the nominotypical
form from Zaire and Uganda. Distribution: Uganda.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
APPENDIX C
Autapomorphies of the species and subspecies of
Savtshenkia
aberdareica: no autapomorphy recognized.
aberdareica ulugurica: no autapomorphy recog-
nized (known of female holotype only).
akeleyi: inner gonostylus with dorsal margin anteri-
or part undulating.
alpha: posterior rim of inner gonostylus ventrally
blackish sclerotized (fig. 161, arrow).
alpium: shape of posterior margin of male tergite 9-
10; shape of posterior margin of male sternite 8 (fig.
105); shape of extension of midventral area (long; fig.
59)
asbolodes inner gonostylus with anterior part rath-
er slender.
aspromontensis shape of inner gonostylus (crest,
fig. 61); shape of male tergite 9-10 (two pairs of
pointed extensions, fig. 135).
aster. shape of aedeagal guide; gonapophysis slen-
der, downcurved and pointed; shape of male tergite
9-10; shape of posterior margin of male sternite 8
(fig. 109).
atlas. extensions of male tergite 9-10 mediolaterally
compressed.
baltistanica: arrangement of setae on posterior mar-
gin of male sternite 8.
benesignata: shape of inner gonostylus (fig. 72).
boreosignata: broad membranous zone of tergite 9-
10 (fig. 138); reduction of spines on lobes on posteri-
or margin male sternite 8; fragmentum with low an-
terior part; shape of aedeagal complex (fig. 99).
breviantennata: anterior part of inner gonostylus
widening towards apex, lateral ridge of inner gonosty-
lus short and angled.
caligo: shape of posterior margin of male tergite 9-
10.
cheethami: midventral area posteriorly with long
conical extension (fig. 35).
chrysocephala: anterior part of inner gonostylus
constricted before tip (lateral view).
confusa: inner gonostylus with two blackish sclero-
tized dentate ridges on lateral side of posterior part
(fig. 58); male tergite 9-10 with truncate extensions
close together; shape of posterior margin of male ster-
nite 8.
corsosignata: shape of inner (fig. 69) and outer go-
nostyli.
cyrnosardensis. inner gonostylus caudodorsally with
posteriorly produced pubescent lobe; outer gonosty-
lus with slender extension on posterior margin at
about two-thirds from base; apex of elongate frag-
mentum medially curved.
draconis. shape of inner gonostylus; reduced nasus.
eleonorae. no autapomorphy recognized.
322
elgonensis no autapomorphy recognized.
eugeni: extensions of male tergite 9-10 upcurved
(lateral view); shape of inner gonostylus; anterior ap-
odemes sperm pump elongate and dorsally directed,
compressor apodemes mediolaterally flattened in api-
cal part.
fragilina: inner gonostylus with acute sclerotized
extension at ventral tip of posterior ridge.
fragilis. aedeagal guide elongate and slender (fig.
22); suture x broad, sigma-shaped.
gimmerthali: inner gonostylus with anterior part
gradually sloping towards bifid apex (fig. 66).
gimmerthali mattheyi: outer gonostylus without
posterior sclerotized appendage (present in both oth-
er subspecies).
gimmerthali pteromaculata: inner gonostylus with
relatively broad anterior part (fig. 67); outer gonosty-
lus broadened above midlength (fig. 84).
glaucocinerea: shape of extensions of tergite 9-10.
goriziensis shape of inner gonostylus (fig. 54).
graciae. posterior margin of male sternite 8 medial-
ly with rounded extension set with long and strong
setae.
grisescens. extensions of male tergite 9-10 dorsoven-
trally flattened (figs. 121, 122).
haennii: fragmentum 3-lobed (fig. 27); inner go-
nostylus with ridge separated in posterior and ventral
part, ventral part ridge extended over posterior part of
inner gonostylus, and dorsal crest anteriorly abruptly
terminating (fig. 68).
hancocki: male antenna reduced in length.
hartigiana: shape of inner gonostylus (long anterior
part, strongly protruding lateral ridge); extension of
midventral area gonocoxites (fig. 33).
holoptera: posterior margin of male sternite 8 with
a few strong setae (fig. 107).
ignobilis shape of outer gonostylus; shape of sp2.
imperfecta: wings reduced in both sexes.
interserta: shape of inner gonostylus; outer gonos-
tylus large, posteriorly inclined.
invenusta: shape of posterior margin of male tergite
9-10 (fig. 136).
invenusta microinvenusta: inner gonostylus anteri-
orly more curved forward than in both other forms;
differing primarily in smaller size and darker colour
from both other forms.
invenusta subinvenusta: inner gonostylus relatively
slender.
jeekeli: shape of inner gonostylus; shape of outer
gonostylus (fig. 80); shape of posterior margin of
male sternite 8; sperm pump with sclerotized sperm
duct; apex of aedeagus with spines.
kiushiuensis: shape of posterior margin of male ster-
nite 8.
koreana: shape of posterior margin of male sternite
8.
letifera: no autapomorphy recognized (probably
very similar to sordidipes and venerabilis).
limbata: anterior part of inner gonostylus produced
into long and bifid extension (fig. 64).
lundbladi: shape of inner gonostylus (elongate, with
short crest); shape of outer gonostylus (fig. 78); shape
of posterior margin of sternite 8 (fig. 103); surface of
sp2 anterodorsally with cone-shaped extension.
macaronesica: shape of extension of midventral area
(short).
mannbheimsi. outer gonostylus with incision ventral
of posterior sclerotized spur.
minuscula: no autapomorphy recognized (probably
very similar to zgnobilis).
mohriana: gonapophysis as slender curved spine.
multipicta: shape of inner gonostylus (high anterior
part with long and slender anterior apex); shape of
outer gonostylus (oblong).
nephrotomoides. blackish species.
nephrotomoides invariegata: no autapomorphy rec-
ognized (known of female holotype only).
nielseni: shape of inner gonostylus; shape of outer
gonostylus (fig. 85); shape of medial extension of
male sternite 8.
nivalis. no autapomorphy recognized (probably
very similar to glaucocinerea).
obsoleta: shape of inner and outer gonostyli; shape
of posterior margin of male sternite 8 (fig. 106); spi-
nous extensions on posterodorsal part fragmentum
Grea):
odontostyla: shape of male tergite 9-10; inner go-
nostylus dorsally with long and dense pubescence
(fig. 63); apical part of fragmentum bent and posteri-
orly directed; gonapophysis twisted.
omega: inner gonostylus with hump in anterior half
of posterior part (fig. 181).
ornata: wing cell a2 almost symmetrical in distal
tip; posterior extensions male tergite 9-10 dorsoven-
trally flattened, laterally curved, ventrally carrying
long thin spinous setae; shape of posterior margin of
male sternite 8; midventral anterior extension of male
sternite 9 elongate, membranous; shape of appendage
of midventral area gonocoxites (fig. 40); suture x
broad, enclosing isolated sclerite; female tergite 9 me-
dially reduced to strip.
pagana: female brachypterous.
pechlaneri: male tergite 9-10 with pair of low extra
extensions lateral of usual extensions on posterior
margin; shape of inner gonostylus (fig. 65).
persignata: posterior part of inner gonostylus corru-
gated (fig. 73).
persignata tofina: differing in details of male termi-
nalia from nominotypical form.
phoroctenia: shape of male tergite 9-10; shape of
posterior margin of male sternite 8; lateral ridge of in-
ner gonostylus posterodorsally angular; suture x of
H. DE Jone: Phylogeny of Tipula (Savtshenkia)
gonocoxite Y-shaped; shape of extension of midven-
tral area gonocoxites (fig. 41).
postposita: shape of outer gonostylus.
productella: no autapomorphy recognized.
rufina: extensions of tergite 9-10 extremely long
and slender; shape of inner gonostylus; tip of gona-
pophysis medially curved; posterior margin of male
sternite 8 bulbously curved.
rufina maderensis. larger than nominotypical spe-
cies.
sardosignata: shape of inner gonostylus; shape of
outer gonostylus (fig. 88); shape of posterior margin
of male sternite 8.
sciadoptera: shape of inner gonostylus.
serrulifera: shape of inner gonostylus (fig. 60);
shape of outer gonostylus; shape of posterior margin
of male sternite 8.
signata: posterior part of inner gonostylus anterior-
ly rounded (fig. 74); midventral extension on posteri-
or margin male sternite 8 large, with dorsal sclerite
(fig. 116).
simulans: inner gonostylus with lateral ridge dorsal-
ly produced into pointed extension; shape of exten-
sion of midventral area.
sordidipes. legs extensively blackened.
staegeri. extensions of male tergite 9 widely separ-
ate, mediolaterally flattened; male sternite 8 midven-
trally conical; posterior extensions of male sternite 8
elongate; aedeagal guide posteriorly curved; shape
and length of gonapophysis; shape of inner gonosty-
lus; female sternite 8 dorsally extended; base of hypo-
gynial valve laterally bulging.
subalpium: shape of posterior margin of male ster-
nite 8.
subnodicornis. tergite 9-10 with extension in
between lateral pair (fig. 128).
subsignata: together with next subspecies distin-
guished by lateral ridge being acutely bent and anteri-
orly produced in ventral part (fig. 70); differing from
subsignata cazorla in details of inner gonostylus.
subsignata cazorla: differing from nominotypical
subspecies by shape of inner gonostylus (anterior part
dorsally abruptly narrowed, lateral view).
subvafra: shape of male tergite 9-10; shape of inner
gonostylus.
tetragramma: wing pattern.
trinacria. outer gonostylus with extremely long
posterior sclerotized appendage (fig. 192); sclerotiza-
tion of proctiger well demarcated (fig. 188).
tulipa: inner gonostylus with relatively short lateral
ridge; midventral area gonocoxites broad and long
pubescent.
venerabilis no autapomorphy recognized (very
similar to sordidipes).
villeneuvii: small and posteriorly curved outer go-
nostylus; gonocoxite posterodorsally acutely angled.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
324
J. KJÆRANDSEN
Museum of Zoology, University of Bergen, Norway
EE INE WESPECIES OF PSEUDEXECHIA
TUOMIKOSKI FROM TANZANIA AND THAILAND
(WIMMERA MYCE VORAILIDAE)
Kjærandsen, J., 1994. Three new species of Pseudexechia Tuomikoski from Tanzania and
Thailand (Diptera: Mycetophilidae). — Tijdschrift voor Entomologie 137: 325-330, figs. 1-4.
[rss 0040-7496]. Published 15 December 1994.
Pseudexechia longistylus sp. n. and P. lanceostylus sp. n. from Tanzania and P. inthanonensis sp.
n. from Thailand are described, based on adult males. The terminalia are figured. The presen-
ce of a pair of medium sized bristle-like dorsocentral setae on scutum in the two African species
and an aberrant wing venation in P. inthanonensis sp.n. is commented on.
J. Kjærandsen, Museum of Zoology, University of Bergen, Muséplass 3, N-5007 Bergen,
Norway.
Key words. — Diptera; Mycetophilidae; Pseudexechia; Tanzania; Thailand; taxonomy.
Pseudexechia Tuomikoski, 1966 constitutes one of
14 genera in the tribus Exechiini, subfamily
Mycetophilinae (Tuomikoski 1966). The genus is
characterised by the absence of large discal setae on
the mesoscutum, by the ovate clypeus and on charac-
ters in the male terminalia such as the bud-like stern-
al process. Chandler (1978) treated all eight holarctic
species known at that time. Three palaearctic species
were added later; viz. P. trilobata Ostroverkhova,
1979, P. ussurensis Zaitzev, 1982 and P. altaica
Zaitzev, 1988. Matile (1970) revised the Afrotropical
Pseudexechia comprising four species. A fifth species,
P. tanganyikae (Lindner, 1958) originally described as
Exechia, was later referred to this genus as well
(Matile 1980). No species of Pseudexechia are hither-
to known from the oriental region. Thus, 16 species
of the genus Pseudexechia were known: 9 palaearctic,
2 nearctic and 5 afrotropical. Of only one of the five
Afrotropical species both sexes were described, the
four other were all described on single females.
Female holotypes make it difficult to describe new
species from the region as certain association of the
sexes is not easy to establish without large samples or
reared specimens. However, based on differences in
external characters as coloration and wing venation
the present specimens can not belong to any of the
described species.
MATERIAL
The material consists of two males collected in the
West Usambara Mountains in Tanzania in 1990, and
one male collected in the Doi Inthanon mountain in
Thailand in 1991. The holotypes are deposited in the
Museum of Zoology, Bergen (ZMBN).
METHODS AND TERMINOLOGY
The specimens were cleared and slide mounted in
Canada balsam. The general terminology follows
McAlpine (1981).
Wing measurements (fig. 1): Total wing length is
measured from the extreme base of the distal median
plate. A = distance between the points where R, and
R,.; reach wing margin. B = distance between the
points where R,, and M, reach wing margin. C =
width of M-fork at wing margin. D = width of Cu-
fork at wing margin. E = distance from extreme base
of the distal median plate to base of M-fork. F = dis-
tance from extreme base of the distal median plate to
base of Cu-fork. Other measurements are self-expla-
natory and indicated in Fig. 1.
The tibial spur formula is the length of each spur in
relation to the apical diameter of tibia in the follo-
wing order: fore tibial spur; the two mid tibial spurs;
the two hind tibial spurs.
SYSTEMATIC PART
Pseudexechia longistylus sp. n.
(figs. 1 - 2)
Type material. — Holotype male, TANZANIA: Tanga re-
gion, W. Usambara Mts, Mazumbai, 1440 m asl,
22.X1.1990, G. E. E. Soli, sweep net (ZMBN No. 179).
Diagnostic characters. — The male imago is charac-
925
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
WING LENGTH —
Fig. 1. Pseudexechia longistylus sp. n., right wing showing measuring points. — Total wing length is measured from the extre-
me base of the distal median plate to tip of wing. A = distance between the points where R, and R,,, reaches wing margin. B
= distance between the points where R, and M, reaches wing margin. C = width of M-fork at wing margin. D = witdh of
Cu-fork at wing margin. E = distance from extreme base of the distal median plate to point of M furcation. F = distance fr-
om extreme base of the distal median plate to point of Cu furcation. M-p. = M-petiole.
Fig. 2. Pseudexechia longi-
stylus sp. n., male termina-
lia. — A, ventral view; B, in-
ternal face of the
gonostylus. Scale 0.50 mm.
terised by the enlarged terminalia in which the dorsal
lobe of gonostylus ends in a very elongated lobe with
a setose cushion basally, and by the shape of the bi-
furcate internal process. The ventral lobe of the gono-
stylus also is enlarged compared to other species.
Etymology. — From Latin, longus, long, and stylus,
style, referring to the elongated gonostylus. A noun in
apposition.
Description of male holotype. — Length of thorax
0.86 mm. Wing length 2.62 mm. Wing length /
length of thorax 3.1. Wing length / length of fore fe-
mur 3.1.
Coloration. Antenna with scape, pedicel and basal
part of flagellum yellow, gradually becoming darker
and more greyish dusted towards tip. Head yellow.
Maxillary palp greyish yellow. Thorax mainly yellow;
scutum with narrow pale yellow or somewhat silvery
shining stripe along lateral margin, delimited by dar-
ker margins below and above; scutellum darker yel-
low. Wing unmarked, yellow tinted, paler towards tip
and hind margin. Wing veins yellow with dark setae.
Halter greyish yellow. Legs mainly yellow; coxae paler
yellow, hind coxa posteriorly with darker stripe.
Abdominal tergites mainly yellowish brown; tergite 1
and 2 with pale lateral markings; tergite 3 to 5 with
pale posterolateral markings, almost reaching middle;
tergite 6 almost entirely dark. Terminalia mainly yel-
low, darker apically on gonocoxite, on sternal process
and on parts of dorsal lobe of gonostylus.
Head. Total length of flagellum 1.35 mm. First fla-
gellomere 1.5 times as long as second flagellomere.
Second flagellomere 2.2 times as long as wide. Five
large orbital setae. Small round median ocellus pre-
sent. Diameter of lateral ocellus / diameter of median
ocellus 1.7. Clypeus bearing 31 setae; width / length
0.8. Fourth palpomere about 1.5 times as long as
third palpomere.
Thorax. Pronotum with vertical row of 3 erect
setae. Scutum with pair of medium sized bristle-like
dorsocentral setae posteromedially in front of scutel-
lum (prescutellar setae); anterior and lateral margin
with large setae; otherwise uniformly clothed with
small setae (but larger and fewer than in other
Pseudexechia except in P. lanceostylus sp. n.).
Scutellum with pair of strong scutellar setae and pair
of very short incurved setae basad of them.
Proepisternum with 1 strong, 1 medium sized and 5
small setae. Laterotergite with 6-7 large and about 50
small setae.
Wing (fig. 1). Crossvein h with 1 ventral seta. R,
with 11 ventral setae apically. R,,, with 26-27 ventral
setae apically. Wing length / length of R, 2.6. Wing
length / length of R,,, 1.9. R,,, nearly straight. Length
of r-m / length of M-petiole 1.7. A/ B 1.4. E / length
KJAERANDSEN: New species of Pseudexechia
of M, 0.7. E / length of M, 0.8. Base of Cu-fork well
beyond base of M-fork. M-fork veins distaly diver-
gent. Cu-fork veins divergent. F / E 1.3. F / length of
@uA, LEN length of @uA, 255 Ce Delve
CuP reaching as far as base of Cu-fork. Vein A, weak,
shorter.
Legs. Length of fore basitarsus / length of fore tibia
1.3. Tibial spur formula 3.4; 8.8, 5.6; 4.6, 4.6. Fore
tibia with 1 ad, 36 p and 3 v setae. Mid tibia with 40
a, 5 pd and 11 p setae. Hind tibia with 8 a, 5 pd and
3 p setae. Posterior sensillae placodea on basal part of
tibia2 33353.
Terminalia (fig. 2A, B). Length of gonocoxite 0.60
mm. Sternal process narrow with about 40 tiny setae;
length / width 1.9. Dorsal lobe of gonostylus 0.84
mm long with very elongated lobe, basally with
strongly setose cushion. Sclerotized internal process
bifurcate with both prongs widening apically. Ventral
lobe of gonostylus large, bearing 6 fan-tipped setae
basally. One lobe of the inner lamellate parts of the
gonostylus apically with a slightly curved, spine-like
seta thus forming a hook. Cerci thin and slender, re-
aching to about apical level of gonocoxite, with two
short but strong apical setae (not figured).
Pseudexechia lanceostylus sp. n.
(fig. 3)
Type material. — Holotype male, TANZANIA: Tanga re-
gion, W. Usambara Mts., Mazumbai, 1530 m asl,
02.-03.X1.1990, G. E. E. Soli, Malaise tent (ZMBN No.
180).
Diagnostic characters. — The male imago is charac-
terised by the shape of the dorsal lobe of gonostylus
which ends in a slender lobe, and by the shape of the
bifurcate internal process.
Etymology. — From Latin, /ancea, a small light spe-
ar, and stylus, style, referring to the slender lance-like
dorsal lobe of gonostylus.
Description of male holotype. — Length of thorax
0.98 mm. Wing length 3.02 mm. Wing length /
length of thorax 3.1. Wing length / length of fore fe-
mur 3.1.
Coloration. Antenna with scape, pedicel and basal
part of flagellum yellow, gradually becoming darker
and more greyish dusted towards tip. Head and
maxillary palp yellow. Thorax mainly yellow; scutum
uniformly brownish yellow with pale yellow or so-
mewhat silvery shining stripe along lateral margin,
delimited by darker margins below and above; scutel-
lum brownish yellow with faint paler median stripe.
Wing unmarked, yellow tinted, slightly paler towards
hind margin. Wing veins yellow with dark setae.
Halter pale yellow with three darker yellow stripes.
327
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 137, 1994
Fig. 3. Pseudexechia lanceostylus sp. n. male terminalia. — A, ventral view; B, internal face of the gonostylus. Scale 0.50 mm.
Legs mainly yellow; coxae paler yellow, fore coxa an-
teriorly with darker stripe. Abdominal tergites mainly
yellowish brown; tergite 1 to 3 with pale lateral mar-
kings; tergite 4 and 5 with pale posterolateral mar-
kings, almost reaching middle; tergite 6 almost enti-
rely dark. Terminalia mainly yellow, darker on sternal
process and on parts of dorsal lobe of gonostylus.
Head. Total length of flagellum 1.53 mm. First fla-
gellomere 1.7 times as long as second flagellomere.
Second flagellomere 2.3 times as long as wide. Six lar-
ge orbital setae. Tiny median ocellus present.
Diameter of lateral ocellus / diameter of median ocel-
lus 2.4. Clypeus bearing about 60 setae; width /
length 0.8. Fourth palpomere about 1.5 times as long
as third palpomere.
Thorax. Pronotum with vertical row of 3 erect
setae. Scutum with pair of medium sized bristle like
dorsocentral setae present posteromedially in front of
scutellum (prescutellar setae); anterior and lateral
margins with large setae; otherwise uniformly clothed
with small setae (but larger and fewer than in other
328
Pseudexechia except in P. longistylus sp. n.). Scutellum
with pair of strong scutellar setae and 2 pairs of very
short posteriorly curved setae basad of them.
Proepisternum with 1 strong, 2 medium sized and 8
small setae. Laterotergite with 8-10 large and about
60 small setae.
Wing. Crossvein h with 1-2 ventral setae. R, with
5-8 ventral setae apically. R,,, with 3-4 ventral setae
basally and 36-39 ventral setae apically. Wing length
/ length of R, 2.6. Wing length / length of R,,; 1.8.
R,,, nearly straight. Length of r-m / length of M-pe-
tiole 1.6. A / B 1.5. E/ length of M, 0.7. E / length of
M,0.8. Base of Cu-fork well beyond base of M-fork.
M-fork veins distaly divergent. Cu-fork veins diver-
gent. F / E 1.3. F / length of CuA, 1.5. F / length of
CuA, 2.3. C / D 1.1. Vein CuP reaching as far as ba-
se of Cu-fork. Vein A, strong, shorter.
Legs. Length of fore basitarsus / length of fore tibia
1.8. Tibial spur formula 3.7; 9.8, 7.5; 5.4, 5.3. Fore
tibia with 1 ad, 39 p and 3 v setae. Mid tibia with 41
a, 5 pd and 7 p setae. Hind tibia with 9 a, 5 pd and 4
KJAERANDSEN: New species of Pseudexechia
Fig. 4. Pseudexechia inthanonensis sp. n. male terminalia. — A, ventral view; B, internal face of the gonostylus. Scale 0.50 mm.
p setae. Posterior sensillae placodea on basal part of ti-
bia 3; 4; 3.
Terminalia ( Fig. 3A, B). Length of gonocoxite
0.57 mm. Sternal process rounded with about 55 tiny
setae; length / width 1.4. Dorsal lobe of gonostylus
0.51 mm, ending in slender lobe. Sclerotized internal
process bifurcate, the the prongs pointing in almost
opposite directions. Ventral lobe of gonostylus broad
and rounded, bearing 6 fan-tipped setae. Inner lamel-
late parts of gonostylus with lobe ending in strong
seta—like hook. Cerci thin, not reaching apical level of
gonocoxite, with two strong apical and one strong
subapical setae (not figured).
Pseudexechia inthanonensis sp. n.
(fig. 4).
Type material. — Holotype male, THAILAND: Chiang Mai
province, Doi Inthanon, about 2200 m a.s.l., 12.1V.1991, J.
Kjærandsen, sweep net (ZMBN No. 181).
Diagnostic characters. — The male imago is separa-
ble from other species by the shape of the dorsal lobe
of gonostylus, which has a smooth rounded hook at
the tip and a large internal process with 25 teeth, by
the shape of the sternal process and by the short stem
of the cubital fork.
Etymology. — From Doi Inthanon, a mountain in
northern Thailand. A noun in genitive case.
Description of male holotype. — Length of thorax
1.20 mm. Wing length 3.82 mm. Wing length /
length of thorax 3.2. Wing length / length of fore fe-
mur 3.3.
Coloration. Antenna with scape, pedicel and first
flagellomere for half of its length pale yellow, rest of
flagellum greyish yellow. Head mainly brown, frons
and sides yellowish. Maxillary palp yellow. Pronotum
yellowish brown. Scutum brown with three yellow
stripes, median stripe broadening to fore margin, and
narrow yellow lateral margins. Scutellum brown with
yellowish lateral margins. Pleura yellowish brown.
Wing unmarked, greyish yellow tinted, paler on basal
half posterior of R-stem. Wing veins yellowish brown
with dark setae. Halter pale yellow with three greyish
yellow stripes. Legs mainly yellow; coxae paler yellow,
each coxa with yellowish brown stripe. Abdominal
tergites mainly brown; tergite 2 to 5 with pale poste-
rolateral markings, reaching middle; tergite 6 dark
brown. Terminalia mainly yellow, darker on sternal
process and on parts of dorsal lobe of gonostylus. Tip
of dorsal lobe of gonostylus whitish.
Head. Total length of flagellum 1.84 mm. First fla-
gellomere 1.7 times as long as second flagellomere.
929,
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
Second flagellomere 2.0 times as long as wide. Four
to five large orbital setae. Small median ocellus pre-
sent. Diameter of lateral ocellus / diameter of median
ocellus 2.1. Clypeus bearing about 40 setae; width /
length 0.8. Fourth palpomere about 1.4 times as long
as third palpomere.
Thorax. Pronotum with vertical row of 2 strong
and 2 medium sized upcurved setae. Anterior and la-
teral margin of scutum with setae of different sizes;
scutum otherwise uniformly clothed with small dark
setae. Scutellum with pair of strong scutellar setae and
pair of short posteriorly curved setae basad of them.
Proepisternum with 1 large, 2 medium sized and 24
small setae. Anepisternum with 6 small setae.
Laterotergite with 12-14 large and about 200 tiny
setae.
Wing. Crossvein h without ventral setae. R, with 9
ventral setae apically. R,,, with 4-5 ventral setae basal-
ly and 58-65 ventral setae apically. Wing length /
length of R, 2.3. Wing length / length of R,; 1.7.R,.
distinctly downcurved. Length of r-m / length of M-
petiole 1.0. A / B 1.9. E / length of M, 0.7. E / length
of M, 0.8. Base of Cu-fork not beyond base of
M-fork. M-fork veins only slightly divergent apically.
Cu-fork large, veins divergent. F / E 0.98. F / length
of CuA, 1.2. F / length of CuA, 1.7. C / D 0.9. Vein
CuP reaching well beyond base of Cu-fork. Vein A,
strong, shorter.
Legs. Length of fore basitarsus / length of fore tibia
1.2. Tibial spur formula 3.5; 6.7, 6.4; 4.6, 4.5. Fore
tibia with 6 ad, 45p and 8 v setae. Mid tibia with 49
a, 5 pd and 4 p setae. Hind tibia with 6 a, 5 pd and 8
p setae. Posterior sensillae placodea on basal part of ti-
Dia3:93 3:
Terminalia ( Fig. 4A, B). Length of gonocoxite
0.67 mm. Sternal process narrow without setae;
length / width 1.8. Dorsal lobe of gonostylus 0.67
mm long with smooth rounded hook at tip and rela-
tivly large sclerotized internal process with 25 teeth.
Ventral lobe of gonostylus bearing 4-5 fan-tipped
ventral setae and one strong dorsal setae apically.
Inner lamellate parts of gonostylus with 5 small setae.
Cerci thin and slender, reaching to about apical level
of gonocoxite, without strong setae (not figured).
Discussion
The presence of a pair of medium sized bristle-like
dorsocentral setae posteromedially above scutellum in
Pseudexechia longistylus sp. n. and P. lanceostylus sp. n.
is not in accordance with the generic description.
Tuomikoski (1966) used absence of ‘discal bristles’ as
one of the main diagnostic characters for
Pseudexechia. The other setae on the scutal disc in
these two species are also relatively strong, erect and
bristle-like when compared with other species in the
330
genus, and they are fewer in numbers. Hence, this
character should be used with care. However, other
generic characters clearly place these species within
Pseudexechia.
P. inthanonensis sp. n. is evidently closely related to
the Palaearctic P. trisignata (Edwards, 1913) based on
the structure of the male terminalia. However, P. in-
thanonensis sp. n. is rather different in coloration (yel-
low thoracic stripes on brown ground) and shows an
aberrant and interesting wing venation. The short
Cu-stem places the base of Cu-fork slightly before ba-
se of M-fork (F / E = 0.98). A long Cu-stem (short
Cu-fork) has been used to group the species of
Exechia s.l. (including Exechiopsis and Pseudexechia )
(e.g. Edwards 1925).
These findings strengthen the view of Tuomikoski
(1966) that Pseudexechia is more closely related to
Allodiopsis and Allodia than to Exechta and
Exechiopsis.
ACKNOWLEDGEMENTS
I am indebted to G. E. E. Soli for giving me the op-
portunity to examine the material collected during
ZMBN’s Tanzania expedition in 1990, and for critical-
ly reading the manuscript. The Tanzania expedition
was funded by the Norwegian Research Council
(NAVE).
REFERENCES
Chandler, P. J., 1978. Notes on the Holarctic species of
Pseudexechia Tuomikoski (Diptera: Mycetophilidae),
with the description of a new British species. —
Entomologist’s Record and Journal of Variation 90:
44-51.
Edwards, F. W., 1925. British fungus-gnats (Diptera,
Mycetophilidae). With a revised generic classification of
the family. — Transactions of the Royal Entomological
Society of London 1924: 505-670.
Matile, L., 1970. Pseudexechia de la région éthiopienne
(Dipt. Mycetophilidae). — Bulletin de la Société entomo-
logique de France 75: 209-214.
Matile, L., 1980. 15. Family Mycetophilidae. — In:
Crosskey, R. W., Catalogue of the Diptera of the
Afrotropical region. — British Museum (Natural History),
London., pp. 216-230.
McAlpine, J. F., 1981. Morphology and terminology -
Adults. — In: J. F. McAlpine et al. (eds.), Manual of
Nearctic Diptera. Volume 1. — Research Branch
Agriculture Canada, Monograph no. 27. Ottawa,
Ontario. Pp. 9-63.
Tuomikoski, R. 1966. Generic taxonomy of the Exechiini
(Dipt., Mycetophilidae). — Annales entomologici fennici
32(2): 159-194.
Received: 24 March 1994
Accepted: 8 August 1994
JOHN T. POLHEMUS', NICO NIESER’ & STEVEN L. KEFFER’
“University of Colorado Museum, Englewood, Colorado, © Tiel, The Netherlands &* James
Madison University, Harrisonburg, Virginia
SYNONYMICAL NOTES ON THE NEPA CINEREA
EMNINAEUS COMPLEX (NEPIDAE: HETEROP TERA)
Polhemus, J. T., N. Nieser & S. L. Keffer, 1994. Synonymical notes on the Nepa cinerea
Linnaeus complex (Nepidae: Heteroptera). — Tijdschrift voor Entomologie 137: 331-336.
[issN 0040-7496]. Published 15 December 1994.
The Nepa cinerea Linnaeus complex is distributed over Europe, North Africa, the Middle East
and northern Asia. Within this complex we recognize three species, N. cinerea, N. sardiniensis
Hungerford and N. anophthalma Décu, Gruia, Keffer & Sarbu. The following are considered
to be junior synonyms of N. cinerea Linnaeus, 1758: Nepa rubra Linnaeus, 1758; Nepa scor-
pioaquaticus De Geer, 1773; Nepa cinerea var. minor Puton, 1886, syn. n.; Nepa seurati
Bergevin, 1926, syn. n.; Nepa cinerea var. major Bergevin, 1926, syn. n.; Nepa dollfusi Esaki,
1928, syn. n.; Nepa cinerea var. orientalis Esaki, 1928, syn. n.; Nepa rubra meridionalis Poisson,
1961, syn. n.; Nepa remyi Poisson, 1961, syn. n.; Nepa cinerea poissoni Tamanini, 1973.
Correspondence: Dr. John T. Polhemus, University of Colorado Museum, 3115 S. York St.,
Englewood, Colorado 80110.
Key words. — Heteroptera; Nepidae; Nepa cinerea; synonymy.
The Nepa cinerea Linnaeus complex is a tightly
knit and very widespread species group in the Old
World, distributed throughout Europe, the
Mediterranean region, and as far east as northwest
China and the Amur region of Siberia. Nepa cinerea
was one of the earliest water bugs to be studied
(Hoefnagel 1592, Moufet 1634, Frisch 1728,
Swammerdam 1737-38), and the morphology has
been extensively investigated, with an excellent treat-
ment and summary of earlier studies by Hamilton
(1931). Despite of this wealth of knowledge, the spe-
cies group nomenclature concerning this insect re-
mains unsettled. A definitive synonymical treatment
is needed for the Catalogue of Palearctic Heteroptera
(B. Aukema & C. Rieger, editors) now being prepa-
red for publication.
The early nomenclatural dilemma regarding the
priority and synonymy of Nepa cinerea versus Nepa
rubra Linnaeus, 1758, first addressed in detail by
Esaki (1926), then by Tamanini (1973), was finally
resolved by the ICZN (Opinion 1335/1985) in re-
sponse to a petition by Kerzhner (1981). Tamanini
(1973) has declared the type locality to be Sweden,
because the first specimens were collected there; he ci-
tes inclusion in Linnaeus’ Fauna Svecica (1761) as
evidence for this. Over the years however, a number
of additional taxa have been proposed, as species, sub-
species and varietal forms, that are either synonymous
or very closely allied with N. cinerea. Esaki (1928),
Poisson (1961) and Tamanini (1973) addressed this
species-group problem, however, far from resolving
the status of these species-group taxa, each of these
authors exacerbated the situation by adding at least
one additional variety.
The following is a brief summary of the taxa and
their supposed differences from Nepa cinerea sensu
stricto, with comments on the validity of the charac-
ters used for differentiation.
Nepa cinerea var. minor Puton, 1886 and Nepa ci-
nerea var. major Bergevin, 1926 were established sole-
ly on the basis of size differences, which is not a diffe-
rentiating characteristic, as it varies widely within
single populations. Jaczewski (1934) examined speci-
mens of N. cinerea minor from Algeria, and stated
that ‘they seem to be conspecific with the typical N.
cinerea L.’ We therefore synonymize both varieties.
Bergevin (1926) established Nepa seurati n. sp. fr-
om Tunisia on the basis of a number of somatic diffe-
rences, which he compared, in a table, with Nepa ci-
nerea Var. minor Puton. Esaki (1928) provided a
photograph of the type of sewrati which does indeed
have a slightly different shape than the cinerea speci-
“mens he illustrates, with the lateral abdominal mar-
gins straight along much of the basal part, widening
posteriorly, and abruptly incurved beyond, as noted
by Poisson (1961: 631). Also the fore femur basally is
wider than the N. cinerea specimens illustrated. This
taxon would therefore seem to be separable from
Nepa cinerea cinerea, however in long series taken fr-
om single populations in Morocco (nr. Tangier bor-
331
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
der, 8.VIII.1955, D. R. Lauck, Polhemus Coll.) and
Portugal (nr. Portinas, Riba la Banho, 15.1V.1976,
N. Nieser, Nieser and Polhemus Collns.) all of these
character states are present along with intermediates.
If considered as a valid subspecies, then many of the
populations from north Africa would probably be at-
tributable to this taxon.
Esaki (1928) described Nepa dollfusi from a single
female collected in Morocco. Some of the differences
from N. cinerea he cited may be attributable to alary
polymorphism (see Larsén 1949, 1955), as the flight-
less forms, such as dollfusi, have an altered morpholo-
gy, e. g. straighter hind margin of the pronotum, and
reduced hemelytral membrane. Esaki provided a pho-
tograph of the type of dollfusi which appears to repre-
sent a malformed specimen, quite possibly resulting
from damage during emergence from the last nymp-
hal stage. The specimen also has reduced antennae,
discussed below, that may have also resulted from a
traumatic ecdysis. The red abdominal dorsum noted
for European specimens of N. cinerea in contrast to
dollfusi is unreliable, as it varies from grey brown to
yellowish to pink or red in individual series from
Europe and the Mediterranean region. Theiry (1981)
noted the great variation in size and coloration (of
both ground colour and abdominal tergites) in the
100+ specimens of Nepa collected over a wide range
of elevation in Morocco, all of which he assigned to
N. cinerea, noting that the parameres and antennae of
all forms were of the ‘type cinerea.” Lindberg (1929)
considered N. dollfusi to be only an aberrant speci-
men of N. cinerea.
Nepa cinerea var. orientalis Esaki, 1928 was foun-
ded on the basis of size and slight differences in an-
tennal and head morphology, all unreliable. We have
not been able to study a long series from the Far East
to determine if there is any discernable difference in
the male genitalia from the western European stock,
but for the present the available evidence leads us to
synonymize this form with N. cinerea.
Hungerford (1928) described Nepa sardiniensis
from the island of that name. He separated it from re-
lated taxa by the lack of a prolongation on the second
antennal segment and slender fore femur. These alo-
ne are questionably enough to justify the status of a
separate species for the populations of Sardinia and
Corsica, however Mazza (1971, 1978) has given addi-
tional evidence for separate species status (see below).
Nepa sardiniensis is the only geographically isolated
taxon in the N. cinerea complex, aside from the uni-
que spatially isolated cavernicolous N. anophthalma
Décu et al (in press). Linnavuori (1960) reported this
species from Israel, Transcaspica and Turkestan, but
according to Tamanini (1973) these records refer in-
stead to N. cinerea cinerea.
For the Palearctic region, Stichel (1955) listed as
332
species Nepa seurati Bergevin, Nepa dollfusi Esaki,
Nepa sardiniensis Hungerford, and N. cinerea (as
Nepa rubra), with minor Puton, major Bergevin and
orientalis Esaki clearly designated as forms of the lat-
ter.
Linnavuori (1960) claimed separation of N. cinerea
and N. sardiniensis on the basis of the short second
antennal segment, broader sternite VII, and differen-
tly formed male clasper of the latter. He stated that N.
cinerea does not occur in Palestine. He also stated that
the anterior femora of Nepa seurati are much broader
basally than in N. sardiniensis.
Poisson (1961) gave figures of the antennae, fema-
le subgenital plate and male genitalia of N. dollfusi
Esaki, N. cinerea cinerea (as N. rubra rubra), N. cine-
rea meridionalis (as N. rubra meridionalis), and Nepa
remyi Poisson, 1961 from Morocco. N. remyi was se-
parated on the basis of slight differences in the shapes
of the male subgenital plate (‘opercule génital’), male
paramere, and the base of the fore femur, all of which
have been seen to be somewhat variable within popu-
lations. We consider this variety to fall within the va-
riability of N. cinerea, thus a synonym. Poisson also
gave a key to separate all of the forms and species
known in 1961 (including Nepa apiculata Uhler from
North America and Nepa hoffmanni Esaki from
China), based on differences in the antennae, the re-
spiratory siphon, shape of the anterior femur, overall
body length.
Seidenstücker (1963) investigated the status of N.
dollfusi, N. sardiniensis and N. seurati, relying mainly
on the antennal structure to conclude that Nepa seur-
ati is a good species, and that Nepa sardiniensis
Hungerford, 1928 (June) is a junior synonym of
Nepa dollfusi Esaki, 1928 (April), and occurs in
Turkey as well as Corsica and Sardinia. If
Seidenstiicker’s analysis were accepted, the N. cinerea
complex would include N. anophthalma, N. cinerea,
N. dollfusi (= N. sardiniensis), and N. seurati. Later
authors have not accepted the synonymy of N. sardi-
niensis (see below).
Mazza (1968) studied the variability of the number
of respiratory horns of 5081 eggs of N. cinerea from
Italy. He noted that the modal number of horns do-
cumented by Hinton (1961) for a British population
(7) was different than the modal number (6) for the
Italian population from La Spezia Province. Later he
analyzed many specimens and concluded that the ge-
ographical forms were impossible to delimit, and that
they form a continuum (Mazza 1974). This evidence
suggests clines within N. cinerea.
Mazza (1971) noted constant differences in the
number of respiratory horns (without giving details)
between N. sardiniensis collected on Corsica and N.
cinerea collected in Toscana (Italy), and further stated
that these two populations were reproductively isola-
ted, as cross breeding produced sterile F1 hybrids.
Later he (Mazza, 1978) examined about 1000 speci-
mens of each species, and using several somatic char-
acters in first order equations he showed consistent
separation, therefore N. sardiniensis and N. cinerea are
presently accepted as distinct species.
Tamanini (1973) discussed all of the previously
mentioned forms except Nepa remyi Poisson, 1961,
and proposed yet another subspecies, Nepa cinerea
poissoni Tamanini from the Pyrenees Mountains of
France and Spain, based on differences in pronotal
morphology and antennae. He compared his new
taxon to N. sardiniensis and N. cinerea cinerea. The
differences he cites are not sufficient to separate this
taxon from the latter, and it was synonymized with
the latter by Nieser & Montes (1984). Tamanini
claims that N. sardiniensis is endemic to Corsica and
Sardinia, and states that N. cinerea dollfusi and N. ci-
nerea meridionalis are of ‘uncertain systematic rank.’
For all of Europe and North Africa, Nieser (1978) lis-
ted only Nepa sardiniensis Hungerford and Nepa cine-
rea, and recognized as subspecies of the latter M. c.
poissoni Tamanini and N. c. seurati Bergevin. Later
Nieser & Montes (1984) synonymized N. c. poissoni
with N. cinerea, without comment.
Nepa anophthalma Décu, Gruia, Keffer & Sarbu
(in press) is separable from all other species of the ci-
nerea complex by five characters; 1. A yellow ground
color (vs. dark), 2. Eyes absent (vs. present), 3.
Hemelytra brachypterous, posterior margin sinuate
(vs. macropterous, posterior margin rounded ), 4.
Midlongitudinal groove of mesosternum absent (vs.
present), 5. Paraterga hirsute (vs. glabrous except for
hairs ringing stigmata). This is clearly the most an-
nectant species of the complex, yet the male genitalia
are very similar to the other members of the group. It
is endemic to the Movile cave system in Romania.
As discussed above, most of the characters used by
various authors to separate species-group taxa in the
Nepa cinerea complex are as variable within popula-
tions as they are between populations, thus must be
considered unreliable for the separation of taxa, espe-
cially when dealing with single specimens or small se-
ries. The antennae have been extensively cited as a se-
parating characteristic notwithstanding their
variability. Tamanini (1973) has shown that the an-
tennal morphology is quite variable, even in a single
specimen, thus the antennal morphology is not a
completely reliable specific character in this genus,
although their form may indicate trends, certainly va-
ty between some populations, and seem to have a
norm within certain populations (vide Seidenstiicker,
1963; Tamanini, 1973). Esaki (1928) illustrated the
antennae of a topotypic specimen of N. seurati, the
type of N. dollfusi, and of N. cinerea (no provenance
given). In dollfusi the prolongation of the second seg-
POLHEMUS ET AL: Synonymy of Nepa cinerea
ment is lacking and segments two and three are fused
and short; in this regard it is very similar to N. sardi-
niensis, ın fact so similar that Seidenstücker (1963)
proposed the synonymy of the two taxa. In seurati the
prolongation of the second segment is moderately
long, and segments two and three are separated and
moderately long, very similar to cinerea except in the
latter the prolongation of the second segment is
slightly longer. Tamanini (1973) studied many speci-
mens from various localities, and illustrated the an-
tennal forms most frequently encountered as well as
aberrant forms rarely seen. The range of morphology
within the species essentially covers the spectrum of
differences used to separate the three species group
taxa discussed above, although the ‘normal’ form of
each conforms roughly to those illustrated by Esaki
(1928). Mancini (1936), who tentatively identified
specimens from Libya as N. cinerea minor, questioned
whether the shape of the antennal segments was a va-
lid character.
One of us (SLK) has studied the male genitalic
structure of several populations of the Nepa cinerea
complex from Finland, Morocco, Sardinia, Romania,
and Iran and can find only one significant genitalic
difference in any of the taxa (see below), although the
genitalia of the cinerea complex are considerably dif-
ferent than the genitalia of the apiculata group which
contains Nepa apiculata and Nepa hoffmanni (Keffer
et al. 1990). However, this general lack of male geni-
talic difference should not be taken as evidence that
species or subspecies designations within the cinerea
complex are invalid. Keffer (1991 and personal obser-
vation) has found that male genitalia are often of lit-
tle taxonomic value at the species level in the
Nepidae. For example, in his revision of the New
World waterscorpion genus Curicta, Keffer (1991)
found male genitalia to be diagnostic for the genus
but largely invariant throughout the range of the ge-
nus. Similarly, Keffer (in prep.) has studied a majori-
ty of the species across the family Nepidae and found
that male genitalic characters often have generic or
species group significance but are often of little value
for species determination. One genitalic character,
paramere shape, has been used extensively by two
nepid taxonomists of the recent past, Jose De Carlo
and Raymond Poisson, in their numerous new species
descriptions. Keffer, however, has found that parame-
res may vary either little or else randomly across spe-
cies groups, or even genera, and thus are not general-
ly valid as a species specific character.
Keffer’s observations with Curicta and other nepid
genera are affirmed in the genus Nepa. Male genitalia
do separate the cinerea and apiculata species groups
(Keffer et al. 1990). However, within the cinerea
complex phallic structures, with one exception, are
either invariant, or they vary randomly, across all
335
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
taxa. For example, parameres appear to vary random-
ly and are therefore of no value in making species and
subspecies determinations. Taminini (1973, figs. 61-
78) figured parameres for N. cinerea cinerea, N. cine-
rea poissoni, N. cinerea seurati, and N. sardiniensis. We
have observed N. cinerea cinerea specimens from
Finland with paramere shapes similar to those figured
by Taminini for N. cinerea poissoni and we have ob-
served N. sardiniensis parameres that are similar to
those figured by Taminini for N. cinerea cinerea.
Further, Décu et al. (in press) have shown that N.
anophthalma has parameres similar to those of N. ci-
nerea poissoni.
Only one genitalic character, the shape of the se-
condary struts, appears to be of value for separating
some cinerea group taxa. During copulation, the ma-
le’s secondary struts and ventral diverticulum lodge in
the female bursa copulatrix (Larsen 1938, fig. 26; se-
condaryasstzutsen ug Gino ea = VE
Vaginaltasche’; Keffer 1991, p.3) where they may ha-
ve a holdfast and/or stimulatory function. If, in fact,
the struts are stimulating the female during copula-
tion, and thus subject to sexual selection, they could
exhibit species specificity. Taminini (1973: 233) no-
ted that, in ventral view, the secondary struts (‘pro-
cessi a spatola della coniunctiva’) of N. cinerea cinerea
specimens from Sicily and continental Italy converge
toward the midline distally, whereas N. sardiniensis
secondary struts are nearly straight. We would add to
Taminini’s description two of our own observations.
First, converging secondary struts, as in cinerea cine-
rea, are also found in the cave dwelling cinerea group
taxon, N. anophthalma and in apiculata group taxa.
Second, in both N. cinerea cinerea and N. sardiniensis
the secondary struts are dorsoventrally flattened whe-
reas in N. anophthalma and in the apiculata group
taxa the struts are tubular in shape. In sum, it appears
that secondary struts can be used to separate N. sard-
iniensis (secondary struts nearly straight, not conver-
ging distally) and N. anophthalma (secondary struts
tubular, not dorsoventrally flattened) from other ci-
nerea group taxa.
The status of four species group taxa must be deci-
ded; they are N. cinerea, N. dollfusi, N. sardiniensis
and N. seurati. The first question to be settled is the
proposed synonymy of dollfusi and N. sardiniensis, by
Seidenstücker (1963). This synonymy seems to rest
on the strength of the similarity of the antennae of the
two taxa. If this synonymy is accepted, and the modal
antennal form is accepted as a valid discriminating
character, then the range of N. dollfusi, the senior sy-
nonym, is Morocco, Tunisia, Corsica and Sardinia.
Tamanini (1973) rejected this synonymy, however,
and also rejected the citation of N. sardiniensis for
Israel by Linnavuori (1960) and for Turkey,
Transcaspica and Turkestan by various authors, as be-
334
ing narrowly based on antennal form only. In this
contention he is supported by Mazza (1971, 1978),
thus current opinion supports the recognition of N.
sardiniensis as a valid species endemic to Corsica and
Sardinia, with N. dollfusi relegated to synonymy un-
der N. cinerea.
Tamanini (1973) contends that N. cinerea cinerea
occurs only as far south as northern Italy, and that N.
cinerea seurati is distributed from southern Italy
through Sicily to Libyia and Tunisia (we would pos-
sibly add Morocco in part). He shows a gradation of
characters on a latitudinal cline through Italy, which
suggests a single species rather than subspecies. If the
subspecies status of seurati were to be accepted, then
dollfusi should be considered as a synonym with aber-
rent antennae. This would leave N. cinerea cinerea as
a variable subspecies with a range throughout nort-
hern Europe and as far south as Morocco nearest
Spain, across the Balkans, through the Middle East,
and as far east as Siberia.
We here reject the subspecies concept for seurati,
however, as based on insufficient evidence and unre-
liable characters, and synonymize it under cinerea.
Mayr & Ashlock (1991:43) give the following de-
finition of the subspecies: “A subspecies is an aggrega-
te of phenotypically similar populations of a species
inhabiting a geographic subdivision of the range of
that species and differing taxonomically from other
populations of that species’. We accept this defini-
tion, and all of the species group taxa of the N. cine-
rea complex that fail to conform to this definition are
placed in synonymy of N. cinerea below. We have
seen no convincing evidence that the characters upon
which these taxa were founded are anything more
than individual variations within and between popu-
lations, scattered more or less randomly, and without
significant separation as required for subspecies char-
acterization. It is indeed possible that sufficient evi-
dence may be presented in the future to reestablish
one or more of the synonymized taxa as valid subspe-
cies, but for the present such clear evidence is lacking.
We therefore recognize only three species, without
subspecies, in the cinerea complex; Nepa anophthalma
(endemic to the Movile Cave system of Romania), N.
sardiniensis (endemic to Corsica and Sardinia), and
the widespread N. cinerea. The results of our investi-
gations are summarized as follows:
Nepa anophthalma Décu, Gruia, Keffer & Sarbu, in
press.
Nepa anophthalma Décu, Gruia, Keffer & Sarbu, in press.
Holotype: 3, Movile Cave, Romania. (ERSI) [examined].
Distribution. — Europe, Romania, nr. Mangalia,
Movile Cave. Endemic.
Nepa cinerea Linnaeus, 1758
Nepa cinerea Linnaeus 1758: 440. Syntypes, d, 9, Sweden
(type locality restricted to Sweden by Tamanini 1973:
226). (uzıu) [not examined].
Nepa rubra Linnaeus 1758: 440. Holotype, 9, locality unk-
nown. (UZIU) [not examined]. (syn. of Nepa cinerea
Linnaeus 1758; see Opinion 1335/1985).
Nepa scorpioaquaticus De Geer 1773: 361. Syntypes, 5, sex
unknown, type locality unknown. (coll De Geer, NHRS)
[not examined]. Unneccessary new name for N. cinerea L.
(see Retzius 1783: 90).
Nepa cinerarea P. L. S. Müller 1774: 472. Incorrect subse-
quent spelling.
Nepa cinerea var. minor Puton 1886: 8. Syntypes, sex unk-
nown, Tunisia, Oued Tessa, ‘Sidi-Mohamed-ben-Ali’,
Oued Bateha, Oued Eddedj, Gafsa, Gabés. (Depository
unknown) [not examined] (syn. Jaczewski 1934, suspec-
ted). Syn. n..
Nepa seurati Bergevin 1926: 290. Holotype, d, Tunisia,
Kebili. (NHN) [not examined]. Syn. n.
Nepa cinerea var. major Bergevin 1926: 294. Holotype, à,
Morocco, Taza. (Depository unknown; coll. Bergevin;
MNHN?) [not examined]. Syn. n.
Nepa dollfusi Esaki 1928: 434 (April). Holotype, 9,
Morocco, Oued Djenanimès près Khénifra. (MNHN) [not
examined]. (syn. Lindberg 1929: 9; restored by
Seidenstiicker 1963: 315, and named senior synonym of
Nepa sardiniensis, synonymy with sardiniensis rejected by
inference by Nieser 1978: 283). Syn. n.
Nepa cinerea var. orientalis Esaki 1928: 436. Syntypes, 39,
Russia, Amur (BMNH) [not examined]. Syn. n.
Nepa rubra meridionalis Poisson 1961: 635. Syntypes, à , 9,
France, Tunisia, ‘Iberian Peninsula’. (coll. Poisson,
USNM) [examined]. Syn. n.
Nepa remyi Poisson 1961: 636. Syntypes, 1d, 29,
Morocco, region d’Ifrane, Tagbaloute. (coll. Poisson,
USNM) [examined]. Syn. n.
Nepa cinerea poissoni Tamanini 1973: 239. Holotype, d,
France, Pyrenées-Orientales, Collioure. (LTRC) [not exa-
mined]. (syn. Nieser & Montes 1984: 38).
Distribution. — Europe, North Africa, Middle
East, northern Asia.
Note. — Nepa annulipes (non Laporte 1833):
Kolenati 1857: 481 is a misidentification referring to
a Laccotrephes species, not collected in the Caucasus;
see Kiritshenko 1918: 171; see also Esaki 1928: 434.
Nepa sardiniensis Hungerford, 1928
Nepa sardiniensis Hungerford, 1928: 120 (June). Syntypes,
14,19, Italy, Sardinia. (semc) [examined]. (syn. with
dollfusi by Seidenstiicker 1963: 322; considered as valid
species by Mazza 1971: 539, 1978: 85, Tamanini 1973:
241).
Distribution. — Europe (Corsica, Sardinia)
Depositories
Abbreviations (codens) for depositories follow
Arnett, Samuelson & Nishida (1993), with the addi-
tion of: ERSI (Emil Racovità Speleological Institute,
POLHEMUS ET AL: Synonymy of Nepa cinerea
Bucharest); LTRC (Livio Tamanini Collection,
Rovereto, Italy).
ACKNOWLEDGMENTS
We thank the following for helpful comments: I.
Lansbury, Oxford; E. Kanyukova, Vladivostok; I. M.
Kerzhner, St. Petersburg. We are grateful to F. Faraci,
Bardolino and A. Carapezza, Palermo, Italy, for assi-
stance with literature and specimens.
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Received: 15 July 1994
Accepted: 10 August 1994
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Volume 137 1994
Tijdschrift
voor
Entomologie
A journal of systematic and evolutionary
entomology since 1858
©,
NIE
DAI
IN IAS
Yaan O
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. J. van Nieukerken (elected 1986) and J. van Tol (1985)
Co-editors
A. W. M. Mol (1990) and R. T. A. Schouten (1990)
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M. Brancucci (Basel), N. E. Stork (London) and M. R. Wilson (Cardiff).
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Published with index of volume 137 (1994).
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Tijdschrift voor Entomologie
Contents of volume 137
Articles
143
155
161
173
271
325
27
57
331
Asche, M. & M. D. Webb
Review of the southern Palaearctic and palaeotropical leafhopper genus
Hengchunia Vilbaste (Homoptera, Cicadellidae).
Baker, D. B.
A new genus of nomadine bees from North Africa (Hymenoptera: Apoidea,
Anthophoridae).
Boer, A. J. de
The taxonomy and biogeography of the lorida group of the genus Baeturia Stal,
1866 (Homoptera, Tibicinidae).
Boer, A. J. de
Four species added to the Baeturia nasuta group, with notes on taxonomy and
biogeography (Homoptera, Tibicinidae).
Garrison, R. W.
A revision of the New World genus Erpetogomphus Hagen in Selys (Odonata:
Gomphidae).
Jong, H. de
The phylogeny of the subgenus Tipula (Savtshenkia) (Diptera: Tipulidae), with
special reference to the western Mediterranean fauna.
Keffer, S. L.: see Polhemus, J. T.
Kjzrandsen, J.
Three new species of PseudexechiaT uomikoski from Tanzania and Thailand
(Diptera: Mycetophilidae).
Kovac, D.: see R. Rozkosny
Malicky, H.: see Weaver, J. S.
Nieser, N.: see Polhemus, J. T.
Patocka, J.
Die Puppen der Spanner Mitteleuropas (Lepidoptera, Geometridae):
Charakteristik und Bestimmungstabelle der Gattungen.
Polhemus, D. A.: see Polhemus, J. T.
Polhemus, J. T. & D. A. Polhemus
Four new genera of Microveliinae (Heteroptera) from New Guinea.
Polhemus, J. T., N. Nieser & S. L. Keffer
Synonymical notes on the Nepa cinerea Linnaeus complex (Nepidae:
Heteroptera).
87 R. Rozkosny & D. Kovac
Adults and larvae of two Ptecticus Loew from Peninsular Malaysia (Diptera,
Stratiomydiae).
87 Tol, J. van
The Odonata of Sulawesi and adjacent islands. Part 3. The genus Macromia
Rambur (Corduliidae).
95 Weaver, J. S. & H. Malicky
The genus Dipseudopsis Walker from Asia (Trichoptera: Dipseudopsidae).
Webb, M. D.: see Asche, M.
Referees for volume 137
B. Aukema, J. Belle, P. L. Beuk, P. Chandler, J. P. Duffels, C. Dufour, R. de Jong,
C. Hauser, I. M. Kerzhner, C. Michener, A. Neboiss, P. Skou, K. D. P. Wilson,
M. R. Wilson.
Dates of publication
Volume 137 (1), pages 1-142, 15 July 1994
Volume 137 (2), pages 143-336, 15 December 1994
© Nederlandse Entomologische Vereniging, Amsterdam Issn 0040-7496
NEW TAXA DESCRIBED IN TIJDSCHRIFT VOOR
ENTOMOLOGIE, VOLUME 137
DIPTERA
Pseudexechia inthanonensis Kjerandsen ................ 329
Pseudexechia lanceostylus Kjaerandsen … … 327
Pseudexechia longistylus Kjerandsen … … 325
Ptecticus malayensis Rozkosny & Kovac … 79
Tipula (Savtshenkia) alpha de Jong ...................... DI
Tipula (Savtshenkia) omega de Jong..................... 315
Tipula (Savtshenkia) trinacia de Jong................... 317
HETEROPTERA
Aegilipsicola Polhemus & Polhemus ...................... 60
Aegilipsicola rapida Polhemus & Polhemus … … … 61
Neusterinsifer Polhemus & Polhemus. … 67
Neusterinsifer compactus Polhemus & Polhemus ....69
Neusterinsifer cyclops Polhemus & Polhemus.......... 69
Neusterinsifer gladius Polhemus & Polhemus......... 70
Neusterinsifer nabire Polhemus & Polhemus.......... 71
Neusterinsifer sepik Polhemus & Polhemus … … …. 72
Tanyvelia Polhemus & Polhemus … 58
Tanyvelia missim Polhemus & Polhemus............... 58
Tarsovelia Polhemus & Polhemus … … 63
Tarsovelia alta Polhemus & Polhemus................... 64
Tarsovelia arfak Polhemus & Polhemus................. 65
Tarsovelia dani Polhemus & Polhemus.................. 65
HOMOPTERA
Baeturia bemmeleni de Boer … … … 10
Baeturia daviesi de Boer ……… … 17
Bacunanonunide Boe inno 20
BACEUTUANCIOUCTOSH AC BOT in one 169
Baeturia hamiltoni de Boer … … … 8
Baeturia hartonoi de Boer … … … nnn 19
Be pia MI AENBOER KI ensen eneen 15
Baeturia retracta de Boer … … i 170
Baeturia silveri de Boer … ……… nnn 13
BaetwnasplendidadelBoers eneen 167
BZB OND EVOH A ACID OCR ennen 13
Hengchunia helleri Asche & Webb … … … … … …. 147
Hengchunia indica Asche & Webb … … … … … …. 150
Hengchunia javana Asche & Webb...................... 152
Hengchunia pakistanica Asche & Webb … … … … 150
HYMENOPTERA
AT D AIO ALES Baker... na 155
Aethammobates prionogaster Baker........................ 156
ODONATA
Erpetogomphus agkistrodon Garrison..................... 199
Erpetogomphus bothrops Garrison … enn 212
Erpetogomphus elaphe Garrison … nennen 205
Erpetogomphus heterodon Garrison … … 230
Erpetogomphus leptophis Garrison … … nn 204
Erpetogomphus liopeltis Garrison ........... 210
Macromia celebica van Tol........... i 88
TRICHOPTERA
Dipseudopsis adiaturix Weaver & Malicky … … … 102
Dipseudopsis flinti Weaver & Malicky.................. 116
Dipseudopsis lucasi Weaver & Malicky................. 123
Dipseudopsis malaisei Weaver & Malicky … … … 124
Dipseudopsis martinovi Weaver & Malicky … … 124
Dipseudopsis robustior andamanensis
Weaver: Ga Mali choy RR RR EA 134
Dipseudopsis schmidi Weaver & Malicky.............. 134
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 137, 1994
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vii
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Tijdschrift voor Entomologie
Volume 137, no. 2
Articles
143 M. Asche & M. D. Webb
Review of the southern Palaearctic and palaeotropical leafhopper genus
Hengchunia Vilbaste (Homoptera, Cicadellidae).
155 D. B. Baker
A new genus of nomadine bees from North Africa (Hymenoptera: Apoidea,
Anthophoridae).
161 A. J. de Boer
Four species added to the Baeturia nasuta group, with notes on taxonomy and
biogeography (Homoptera, Tibicinidae).
173 R. W. Garrison
A revision of the New World genus Erpetogomphus Hagen in Selys (Odonata:
Gomphidae).
271 H. de Jong
The phylogeny of the subgenus Tipula (Savtshenkia) (Diptera: Tipulidae), with
special reference to the western Mediterranean fauna.
325 J. Kjærandsen
Three new species of PseudexechiaTuomikoski from Tanzania and Thailand
(Diptera: Mycetophilidae).
331 J. T. Polhemus, N. Nieser & S. L. Keffer
Synonymical notes on the Nepa cinerea Linnaeus complex (Nepidae:
Heteroptera).
© Nederlandse Entomologische Vereniging, Amsterdam
Published 15 December 1994 ISSN 0040-7496
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