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HARVARD UNIVERSITY
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Comparative Zoology

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DEEL 120 1977

TIJDSCHRIFT
VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

Tijdschrift voor Entomologie, deel 120, 1977

NEDERLANDSE ENTOMOLOGISCHE VERENIGING

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The journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea.
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Afleveringen 1-2 verschenen 15.11.1977
Afleveringen 3-5 verschenen l.vii.1977
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ISSN 0040-7496

INHOUD VAN DEEL 120

Deltshev, C., zie Helsdingen, P. J. van

Helsdingen, P. J. van, K. Thaler & C. Deltshev. — The tenuis group of Lepthyphantes Menge
(Acaneaenbinyphiidae) iii ieee ee ae ee ee Ne ee

Krikken, J. — Asian bolboceratine scarabs of the genus Bolbogonium Boucomont (Coleoptera:
Geotrupidae) Bei ee en Re eN

Nieser, N., zie Vepsäläinen, K.

Roskam, J. C. — Biosystematics of insects living in female birch catkins. I. Gall midges of the
genus Semudobia Kieffer (Diptera, Cecidomyiidae) ........................

Thaler, K., zie Helsdingen, P. J. van

Vecht, J. van der. — Studies of Oriental Stenogastrinae (Hymenoptera Vespoidea) ...........

Vepsäläinen, K., & N. Nieser. — Life cycles and alary morphs of some Dutch Gerris species
(Heteroptera @ernidae)e ar. a reas a en I

Willemse, F. — A study on the genus Cranaella Ramme (Orthoptera, Acridoidea, Catantopinae)

Willemse, F. — A study on the genus Cranae Stal (Orthoptera, Acridoidea, Catantopinae)

Willemse, F. — Classification and distribution of the Sexavae of the Melanesian Subregion
(Orthoptera, Tettigonioidea, Mecopodinae) ..............................

RÉSISTER ne ne A RA I LA RR SEO Ri

DEEL 120 AFLEVERING I 1977

ze mus. COMP. ZOOË
LIBRARY

MAR 2 3 1977

TIJDSCHRIFT nass

V O O R E NTO M O IE O CGIE

_ UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

P. J. VAN HELSDINGEN, K. THALER and C. DELTSHEV. — The tenuis group of
Lepthyphantes Menge (Araneae, Linyphiidae), p. 1—54, fig. 1—69.

Tijdschrift voor Entomologie, deel 120, afl. 1 Gepubliceerd 15-11-1977

THE TENUIS GROUP OF LEPTHYPHANTES MENGE
(ARANEAE, LINYPHIIDAE)

by
P. J. VAN HELSDINGEN

Rijksmuseum van Natuurlijke Historie, Leiden

K. THALER

Zoologisches Institut der Universitat, Innsbruck
and

C. DELTSHEV
Zoological Institute of the B.A.N., Sofia

With 69 text-figures

INTRODUCTION

Within the Linyphiidae, the genus Lepthyphantes is one of the largest. It is rich in
species, which are found in a wide range of habitats, from the Arctic in the north to
the Mediterranean in the south, at high altitudes in the Alps as well as in the lower
parts of Europe. In Asia, North America, and Africa they occur in comparable
situations. A number of species have also been described from other parts of the
world, but at least some of these we may regard sceptically. Even with this
restriction there is an overwhelming amount of species and the genus is notorious
for that very reason. The inaccessibility of the genus is aggravated by the many
poor descriptions without, or with inadequate, illustrations.

Several authors have tried to arrange the species into natural, monophyletic
groups. One of these groups, the tenuis group, will be discussed in this paper. As
already pointed out by Wanless (1973: 139), this group is not sharply limited.
Notably species as L. alacris (Blackwall) and its near relatives are, at least
morphologically, close to the tenuis group, but for practical reasons we have
restricted ourselves to the species-group as delimited by Simon (1929, “4e
Groupe”, p. 589), Locket & Millidge (1953, “Group III”, p. 384), and Wiehle
(1956, “IV. Tenuis-Gruppe”, p. 191). We have included as many species as possible
which from their descriptions, and above all from the illustrations, could be
referred to this species-group. Inevitably some may have escaped attention,
because the available descriptions lacked any indication of their belonging to the
group dealt with here.

Many species of the tenuis group are very common and widely distributed, and
every ecologist will collect large series of certain species in his pitfalls or through
hand-collecting. Still the identification even of the common species is far from
easy, because the palps and epigynes are very small, while the characters

I

ho

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

commonly used for recognition of the species are found in the shapes of the
different elements of these organs. The shapes vary with the angle of vision and
even a slight expansion of the haematodocha, as it is sometimes found in preserved
specimens, may present a very different picture. As a rule, the major collections
examined in recent years contained many a misidentification, a clear demonstra-
tion of the difficulties presented by this group. It also compels us to make a very
critical use of literature records.

It was thought to be helpful to give a summary of all known species of this
group, presenting also their diagnostic characters, their geographical variations,
their distributions and ecological requirements (where possible). Wanless (1971)
already published a series of excellent illustrations, photographic pictures and line-
drawings, of the epigynes of the West-European species. They are of great help and
one realizes how badly similar figures of the male palps are needed. If all former
descriptions had been furnished with such illustrations, a revision of Lepthyphantes
would have been a less time-consuming task.

The main object being to expound our modern views concerning this group, we
nevertheless have also tried to bring the nomenclature up to date. The present
situation is far from satisfactory in view of the fact that many of the older names
were listed as synonyms by authors such as Blackwall, Thorell and Pickard-
Cambridge. Apparently they were in contact with each other from time to time, as
Thorell, for instance, actually was able to compare many examples of Blackwall’s
species with his own specimens. This often resulted in the recognition of
synonymies, and from then onward a name could be unanimously treated as such.
The difficulty lies in the supposition, or near certainty, that recognition of species
was a matter of general appearance rather than of a study of the genitalia,
stridulating files and other structural characters we now consider of paramount
importance. This latter method started with Blackwall, Kulczynski, and Simon,
but we still have to deal with an inheritance of species described with the use of
coloration and abdominal patterns only and listed as junior synonyms of currently
used names. It is beyond our possibilities to evaluate the earlier decisions, given
mainly by Kulczynski and Chyzer & Kulczynski. However, a re-examination of
some of the old collections gives one the strong impression that the recognition of
many of the Lepthyphantes species proved to be difficult even to the more
advanced arachnologists. Too many series were found to consist of more than one
species. It makes it the more important to stabilize the present situation by
procuring useful means for the identification of the species.

THE TENUIS GROUP

Leptyphantes, 4e Groupe, Simon, 1929: 589.
Lepthyphantes, Group III, Locket & Millidge, 1953: 384.
Lepthyphantes, IV. Tenuis-Gruppe, Wiehle, 1956: 191.
Lepthyphantes, Group 1(a), Wanless, 1973: 139.

Within the large and rather heterogeneous genus Lepthyphantes the tenuis group
has been recognized by most authors as a distinct group. It can be easily
distinguished from most of the other currently recognized species-groups, though

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 3

Wanless (1973: 139) indicated the weak points in the system of grouping. He
suggested a much finer subdivision of the genus into species-groups and
subgroups, at the same time pointing out the transitions from one group to the
other. In his grouping of the species — working with the British species and basing
his conclusions on a study of the female genitalia only — all species usually
referred to the tenuis group are kept together in his group I(a), while L. alacris
(Blackwall) (group 1 (b)), and L. leprosus (Ohlert) and L. minutus (Blackwall)
(group | (c)), are thought to be very closely related to the tenuis group proper.
These three species have an epigyne comparable in structure to that of tenuis, but
it is larger and more complicated. It is our present opinion that cristatus takes a
rather isolated position within the tenuis group and shows an affiliation with the
three species placed near the tenuis group by Wanless.

Though recognized by most authors as a homogeneous species-group, it is not
easy to delimit the group clearly. Our diagnosis is given below. Without a detailed
study of the other species-groups it cannot but have its imperfections.

Diagnosis of the tenuis group. Comparisons with other species-groups are given
in square brackets.

Medium-sized animals (1.7—4.1 mm) in comparison with the other species-
groups of the genus. Abdomen of female with a dorsal pattern of transverse
blackish bars and white blotches on the intervening areas, and with light or white
longitudinal lateral band [cf. pallidus group, where faint bars may be present];
pattern obscure in the male. Chelicerae with three dorsal and three ventral teeth,
slightly modified in the male; stridulating files always present. Femora spineless
with the exception of a prolateral spine on femur I. Tibia I with one pro- and one
retrolateral spine, tibia II with one retrolateral spine, in addition to the two dorsal
spines; tibiae III and IV without lateral spines (but with one retrolateral spine in
cristatus); no ventral spines present [cf. L. expunctus, mughi, etc.]. Metatarsi with a
single dorsal spine (but with several spines in spiniger) (cf. nebulosus group, where
several spines are present]. Tm I (0.15—) 0.20—0.25 (—0.30) [cf. obscurus group].
Male palpal tibia with one spine (or more: cristatus), which is at least as strong and
long as the patellar spine [cf. obscurus, pallidus, expunctus, mughi, and related
species]. Cymbium rather simple, without tubercles or horns (but with small
tubercle in cristatus) [cf. obscurus group, nebulosus group]. Epigyne moderately
large, not much protruding from the ventral surface of the abdomen [cf. nebulosus
group]. Scape simple-folded [cf. alacris, where it is invaginated, see Wanless, 1973:
141, figs. 24D—E].

General structure of the secondary genitalia.

Male palp (Fig. 3, 5). — Cymbium (c) not modified, except in cristatus, where
a basal latero-dorsal projection is present. Paracymbium (pc) relatively simple in
comparison with other species-groups within the genus; distal branch of the
roughly U-shaped element ending in a flat, blade-like tip; in most species one or
more teeth present on basal branch or middle section. Haematodocha and tegular
section without diagnostic features on species level; median apophysis (ma) always
with acute tip, which points outward just below tip of cymbium (generic

4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

character). All other elements attached to each other and to radix by membranes,
forming together the radical complex, which looks rather rigid in its mutual
cohesion. Radix (r) always distinctly curved, other elements implanted on basal
and concave middle part (as oriented in the unexpanded palp). Lamella (/) in most
species S-shaped, membranous or lightly sclerotized, free tip usually incised,
forked, or composed of several branches; element conspicuous in lateral aspect.
Embolus (e) a sinuous element which broadens out halfway into an often
dentigerous protrusion, then, after a narrow middle section, widens into a more
voluminous apical section, which is bifid, one of the tips being the spermduct-
tooth. Terminal apophysis (ta) consisting of membranous sclerites and chitinous
tooth or teeth, situated between embolus and lamella. Embolic membrane (em)
small, covering tip of embolus in the unexpanded palp.

Main diagnostically important characters are: positions and size of tooth or
teeth on paracymbium; shape of lamella; absence, presence, number, and size of
teeth on basic section of embolus.

Epigyne (Fig. 1, 2). — Essentially not differing from general structure of this
organ in other species-groups of the genus (see Wanless, 1973, for a schematic
representation of different degrees of complexity). Atrium surrounded by more or
less unmodified, sclerotized anterior and lateral walls; atrium bordered posteriorly
by the posterior median plate (mp) (subgenital sclerite of Wanless), which in all
species of the tenuis group is deeply incised medially; lateral margins of posterior
median plate in posterior view of the uncleared epigyne indicated by the two
darker (more sclerotized) fertilization-ducts. Atrium for the major part covered by
the scape, which runs from its base on the anterior wall of the atrium in posterior
direction; scape in all species of the tenuis group composed of a more or less
straight basal part, reaching as far as the posterior median plate, then rather
abruptly curving to the dorsal side and forward again (inside, and out of view) and
continuing as a second, arched, inner section, the stretcher (st) on the extreme tip
reaching the posterior side of the epigyne just between transition of basal into
arched section and posterior median plate; stretcher always visible in the ventral
aspect at the posterior tip of the visible scape as a round, protruding knob and
furnished with a pit or socket; ventral aspect of epigyne allowing of a view of the
exposed basal part of the scape, bordered at either side by the lateral wing-like
extensions of the first part of the arched, second section of the scape, which in
most species is distinctly wider than the anterior half of the basal part of the scape,
and often of characteristic shape. In the unexpanded or uncleared epigyne, the tip
of the arched part of the scape remains invisible, with the exception of the
stretcher; on this tip are to be found two lateral sockets (see below) and, slightly
more medially, the entrances of the two spermducts, which run from these pores
symmetrically and often parallel through the whole length of the scape and the
anterior and lateral walls of the atrium, each curving to its own side, to the
receptacula seminis.

Functional aspects. — A general understanding of the functioning of the type
of secondary genitalia found in this species-group may be obtained by comparison

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 5

of the structures found here and in Lepthyphantes leprosus (Ohlert), a representa-
tive of another species-group. The functioning of the genital organs in the latter
species have been studied by one of us (Van Helsdingen, 1965), and though we
should be aware of the dangers that attach to a too hasty generalization, we may
safely assume at least some of the elements to function in a similar way.

The presence in the epigyne of a long, curved scape and a characteristically
built male embolus (with its typical, voluminous, double-tipped and at one side
grooved apical section), most probably correlate with each other: in functional
contact the curved part of the scape is pulled around the apical section of the
embolus in such a way, that one tip of the embolus fits into one of the lateral
sockets at the tip of the scape (when the left palp is used it is pushed into the left
socket, and vice versa), while the other tips, the spermduct-tooth, disappears into
the entrance of the female spermduct at the same side of the scape. The scape is
pulled out and kept in place by means of the median apophysis, the tip of which
element grips the stretcher at the tip of the scape, using the socket at the
stretcher’s knob-shaped tip as a foothold, during the very first moments of contact
between palp and epigyne. Leavingthe movements of the different elements out of
consideration, we may assume to find, in the final situation, the male palp firmly
anchored on the epigyne through the combined forces of the paracymbium
(anchored on the scape), the radix (on the scape), the terminal apophysis (within
the atrium and on the posterior median plate), and the lamella (on the posterior
median plate, in co-operation with the terminal apophysis). Whether the scape is
lifted out of its resting position and twisted sideways, as was observed in
L. leprosus, is questionable, but, though likely, not necessarily the case. It would be
very interesting to learn what variations are possible on the theme found in
leprosus, not only in order to understand the intricate movements of the scape, but
also to find out the possible function of the tooth, or teeth in some species, on the
paracymbium.

KEY TO THE RECOGNIZED PALAEARCTIC SPECIES

Males
1. Palpal tibia with four spines, cymbium with a dorso-lateral tubercle near its
BASALE) a esas ER a cristatus (p. 40)
— Palpal tibia with a single spine, cymbium without a dorso-lateral tubercle near
(CSA ASC HN EEE Ee A n 2
2. Posterior margin of paracymbium with tooth orteeth ............. 3
— Posterior margin of paracymbium without tooth, or, at the most, with a ridge
OEREN REIN go Sri Da Aen ES. Be ertoe Ne à 12
3. Paracymbium with small tooth at posterior margin only (Fig.9) ...... 4

— Posterior margin of paracymbium with a large tooth (Fig. 47); or posterior
margin with two teeth (Fig. 33); or a second tooth present at anterior margin

of basal branch (Fig. 18) or at base of distal branch (Fig. 36) ........ 9
dalsenethtoncephalothorax0!0Smmmionless WEAR eee 3
— Jenethioncephalothorat 0.95 mimonmore. ET M E 7
SSuEmbolusswithoutidentielessenfr ante nn PE re 6

— Embolus with denticles along ventral margin of basal section . perseus (p. 11)

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

. Stridulating files fine (Fig. 55); chelicerae without latero-basal boss; lamella

sickle-shapedi(Migs5) Er. 2.0 RIA A mengei (p.8)
Stridulating files coarse and often obscure (Fig. 56, ©!); chelicerae with
latero-basal boss (Fig. 12); terminal and subterminal parts of lamella more
straight, not sickle-shaped (Fig 9) e921 EERE flavipes (p. 12)

. Metatarsi with one dorsal spine only; basal section of embolus with 7 denticles

OT TESS: a Rn Pa Re ERE 8
Metatarsi with several spines; basal section of embolus with about 11 denticles
(Fig. 29); lamella (Fig. 28) with longest branch forked at tip; stridulating files
veryifine(Fign62) i Rn alae HEE na hee ay ENT PRES spiniger (p. 27)

. Stridulating files very coarse (Fig. 60); basal section of embolus with 4

denticles (Fig. 23); dorsal branch of distal part of lamella bluntly truncated
(RISE) EI We at A SERRES SLA drenskyi (p. 22)
Stridulating files less coarse (Fig, 61); basal section of embolus with 7 denticles
(Fig. 26); small tooth at posterior margin of paracymbium pointing backwards
(Fig 27) ee ERE LOTUS AURA BEREN EM renee a zimmermanni (p. 23)

. Larger specimens (length cephalothorax 1.2 mm or more); tooth at posterior

margin of paracymbium large (Fig. 47); distal branches of lamella diverging

(Fig. 47) stridulating files tine (Hip 67) ame nigriventris (p. 36)
Smaller specimens (length cephalothorax 1.1 mmorless) .......... 10
. Only one tooth present at posterior margin of paracymbium ........ 11
Two teeth at posterior margin of paracymbium (Fig. 33) ... floriana (p.28)

. Large second tooth present on paracymbium just inside distal branch (Fig.

36); lamella large but simply curved (Fig. 36) ........ tenebricola (p. 30)
Small second tooth on paracymbium situated at anterior margin of proximal
branch (Fig. 18); lamella smaller, with bluntly tipped dorsal branch (Fig. 18)

A eea noten aan ed ORE tenuis (p. 17)

. No tooth present on paracymbium (Fig. 20); cephalothorax with a dark

median vittae ee te se N tenebricoloides (p.21)
Paracymbium with distinct tooth, large or small ................ 13

. Paracymbium with a denticle near anterior margin of proximal branch (Fig.

13); specimens small (length cephalothorax 0.9 mm or less); lamella (Fig. 13)

hardlyeurved: LIO ND oe EO RO, ene En herbicola (p. 16)
Paracymbium with a large tooth near base of distal branch; specimens larger
RE 0 IR 14

. Distal branches of lamella parallel (Fig. 45); stridulating files fine (Fig. 66)

EEN LOEB RAI TRS AC SPOED AAE RE RE jacksonoides (p.34)
Distal branches of lamella distinctly diverging (Fig. 39); stridulating files
rather coarse (Fig! OS)? Lan nd. a CAL Rene Rte jacksoni (p.32)

Females

. Tibiae III and IV with an I-spine in addition to the two d-spines; epigyne, Fig.

DAE N alas cristatus (p. 40)
Tibiae II and IV with the usual pair ofd-spinesonly ............. 2

. Visible basal section of scape with a medial pit (Fig. 49, 51), or with a

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 7

depression near the broadened tip (Fig. 42,46) . ................ 3
Visible part of scape without a medial pit, nor with a depressed area near its
(Hijo) MEN A TESTO A LOGE a 6
Mm Basalipartofscape withta medial pit (Rigs 49) TEO RR 4
Basal part of scape with a depressed area near tip (Fig.42) ......... 5

. Larger specimens, with length of cephalothorax more than 1.20 mm, femur I

1.5 mm or longer: two (sub)species which differ only slightly in the shape of
their scapes (Fig. 49 and 51) and in their provenance ................

REN OVER EISEN OUT MOT nigriventris (p. 36) and camtschaticus (p. 38)
Smaller specimens: length cephalothorax less than 1.20 mm, femur I not
longer than 1.30 mm; scape of epigyne with a transverse, pigmented
(sclerotized) ridge at transition of exposed basal part and curved inner part
(Fig. 38); medial pit not always present (see also couplet 8) tenebricola (p. 30)

AStrdulatmenlesrathencoarse (Fig 65) en IE: jacksoni (p.32)

Stridulatnmetleshnen(tig1600) PERRET EE jacksonoides (p.34)

. Visible part of scape gradually widening with concave margins from base to

posterior end, without parallel-sided anterior section, the margins reaching
broadest point of scape without any interruption ................ fl
Visible part of scape roundish or with lateral lobes, not gradually widening
with concave margins, but often with a distinctly parallel-sided anterior
section; or, if gradually widening, then lateral margins not reaching broadest
PointoLseapeswithoutanyzinterzuptione PEN Re OI 9

. Larger specimens: length cephalothorax 1.25—1.55, length femur I 1.50—1.80

mm; cephalothorax with a dark grey median streak, abdomen with paired
black spots, otherwise light; stridulating files rather fine (Fig. 59) ........

nés AN ANR dra Pa Ne nd CRÉAS A N tenebricoloides (p.21)
SMAlerispecimensig, rato LR CU PCA 8

. Scape with distinct, more sclerotized, transverse ridge at posterior margin of

exposed part; most specimens with a trace of a medial depression or pit at half
length of scape, in some specimens this pit distinctly developed (Fig. 37 and
38) (see also couplet 4); posterior median plate not much broader than scape;
lateral lobes of inner part of scape with convex margins; width of scape
OND N ey. cose ere eae IE A ER E tenebricola (p.30)
Scape without distinct transverse ridge; posterior median plate distinctly wider
than scape; lateral lobes with concave margins (Fig. 15); width of scape
PREME Ae i e RO bee à herbicola (p. 16)

. Scape with parallel-sided basal “stem” or gradually widening in posterior

Gin CHO Meese a te thea a tet ei eig 10
Exposed part of scape with lateral lobes near base, or broadly rounded 13

. Scape distinctly anchor-shaped, with narrow, parallel-sided “stem” gradually

widening into transverse posterior part; inner, curved part of scape visible as
strongly rounded lobes; posterior median plate not much wider than scape
(Fig. 16); width of scape 0.13—0.15 mm; stridulating files rather coarse (Fig.
DIG) MR NE CAE ERR Me MEL eee nl, Ne tenuis (p. 17)

Baterallobesinotroundedi mare nn ee ee 11

8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

11. Metatarsi with more than one spine; stridulating files very fine (Fig. 62)
EEG TAMA SRT MUR BN NE PONTS eo spiniger (p. 27)

— Metatarsi with a single d-spine; stridulating files not very fine ....... 12
12. Lateral lobes of inner part of scape with rather even concave margin towards
broadest point, from there curved inward (Fig. 24); scape slightly narrower
than in next species (width of outer scape 0.150—0.175, of inner scape
031500417 Simim) a FREE EN han PRES zimmermanni (p. 23)

— Lateral lobes with a small but distinct projection just in front of broadest
point, hence margin in front of broadest point not evenly concave (Fig. 34);
scape slightly wider than in former species (width of outer scape 0.175—0.190,

Ofinnerscape 0.185 0200: mm) MEE SPE NE floriana (p. 28)
13. Exposed part of scape with lateral, wing-like extensions on anterior half (Fig.
d)astridulatingstilestine (ists) ERRE ESS mengei (p. 8)

— Exposed part of scape more or less evenly rounded, i.e. without incisions and
without distinct, more or less parallel-sided basal “stem” (Fig. 11); stridulating
files vercoarse hardlyavisible (Fig SG) EE n flavipes (p. 12)

Lepthyphantes mengei Kulczynski
(Fig. 3—5, 55)

Lepthyphantes mengei Kulczyúski, 1887: 267, 320, Pl. 7 Fig. 37—39 (descr. 9 3 ; Poland). Miller, 1947:
26, 29, 40, 79, Pl. 12 Fig. 8—11, 14, PI. 13 Fig. 1(Q g ). Locket & Millidge, 1953: 388, Fig. 231 A, C,
E, 232 C, D (9 4 ). Wiehle, 1956: 202, Fig. 340—344 (9 3 ). Wanless, 1971: 22, 26, PI. 2, PI. 6 Fig. F
(9): 1973: 129, 134, PI. 1 Fig. 6, Pl. 6 Fig. 6 (9).

Lepthyphantes gallicus Simon, 1929: 591, 594, 734, Fig. 906—907 (deser. 9 & ; France). Jackson, 1930:
653 (= mengei).

Linyphia concinna L. Koch, 1879: 37, PI. 1 Fig. 26 (descr. 9; Siberia). Preoccupied by Linyphia concinna
Thorell, 1875 [ = Centromerita concinna (Thorell)].

Linyphia tomskica Strand; Ermolajev, 1934: 130 (nom. nov. pro Linyphia concinna L. Koch, 1879).

Linyphia concinnella Roewer, 1942: 578 (nom. nov. pro Linyphia concinna L. Koch, 1879).

Lepthyphantes concinnus; Holm, 1945: 8, 56 (as senior synonym of Lepthyphantes mengei Kulczynski).

Remarks. — Despite the recent discovery (see Locket, Millidge & Van
Helsdingen, 1970: 90) that the type-series of Linyphia tenebricola Wider, 1834,
belongs to the species currently known as Lepthyphantes mengei Kulczynski, 1887,
we maintain the name commonly used before that discovery. A proposal to
stabilize this situation has been presented to the International Commission on
Zoological Nomenclature. See also under remarks on Lepthyphantes tenebricola
(Wider).

Both Strand and Roewer published replacement names for Linyphia concinna L.
Koch, 1879, because it was preoccupied by Linyphia concinna Thorell, 1875. Holm
(1945) re-examined Koch’s material and found it to be synonymous with
Lepthyphantes mengei. However, by placing L. mengei as a junior synonym of
Lepthyphantes concinnus he forgot to deal with the problem of the preoccupation.
Subsequently he corrected this error by placing L. concinna as synonym of mengei,
the only valid solution.

The original material of Linyphia concinna L. Koch (ls 29) is preserved in
Stockholm (NRS). Original material of Lepthyphantes gallicus Simon could not be

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 9

Fig. 1—2. Schematic representation of epigyne in Lepthyphantes. |, ventral aspect; 2, lateral aspect (mp,
posterior median plate; st, stretcher).

Fig. 3—5. Lepthyphantes mengei Kulczynski. 3, radical section of male palp, ventral aspect; 4, epigyne;
5, male palp, lateral aspect (c, cymbium; e, embolus; em, embolic membrane; /, lamella; ma, median
apophysis; pc, paracymbium; r, radix; ta, terminal apophysis). 3, X 440; 4, X 225; 5, X 180

10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

traced with any certainty, but all material in the Simon collection under that name
has been examined (MP). Neglecting the presence of a few specimens of other
species — a not unusual sample pollution of Lepthyphantes in old collections — all
gallicus material belongs to mengei, and Jackson’s (1930) statement can only be
confirmed.

Diagnosis. — L. mengei belongs to the smaller species of the group. The fine
stridulating files distinguish it at once from L. flavipes (Blackwall), the shapes of
the lamella and epigyne are quite characteristic.

Measurements (in mm). Total length, 9 1.9—2.5, 3 1.8—2.2; length cephalo-
thorax, 9 0.80—0.92, x 0.80—0.95. Legs: Fe I 1.15—1.30 times length cephalo-
thorax in 9, 1.20—1.35 ing .

Stridulating files fine (Fig. 55).

Male palp (Fig. 3,5) characterized by the comparatively obtuse (if compared
with flavipes) tooth on the posterior margin of the paracymbium, the sickle-shaped
lamella which is provided with a dorsal lobe, the tongue-shaped structure of the
terminal apophysis as seen in the lateral aspect, and the absence of denticles on
the basal part of the embolus. The tooth on the paracymbium lies rather far to the
ventral side. Length of cymbium 0.28—0.37 mm.

Epigyne (Fig. 4). Basal part of scape roughly anchor-shaped but anterior portion
furnished with lateral wings, which slightly protrude in ventral direction; inner
part of scape visible as a slightly projecting lobe at either side of the scape in the
incisions between tips of anchor and lateral wings; stretcher as usual. Posterior
median plate hardly wider than scape. Width of basal part of scape 0.12—0.15, of
posterior median plate 0.13—1.17 mm.

Distribution. — Bonnet could list the species from most European countries
within the following boundaries: the British Isles, Iceland, Scandinavia, the
European U.S.S.R. and West Siberia, the Balkans, Austria, Switzerland, and
France. It has also been recorded from Greenland (Bonnet, 1957: 2432), but this
was proven to be incorrect (Braendegard, 1958: 87). Recent records, later than
1940, include again Siberia (Holm, 1970), Finland (Lehtinen & Kleemola, 1962;
Huhta, 1965, 1971; Palmgren, 1972), Estland (Vilbaste, 1973), Lapland (Holm,
1945, 1952, 1959), Iceland (Cloudsley-Thompson, 1948; Braendegàrd, 1958), and
most other countries within the boundaries mentioned.

A cave record from Rumania (Denis, 1952: 12) was based on four specimens
(MP, re-examined) which appeared to belong to L. flavipes (Blackwall).

We have studied material from Austria (Tirol, Osttirol, common), Czechoslo-
vakia (Böhmen, common), Switzerland (Schwyz, Glarus, Engadin), northern Italy
(Bergamo, Lago di Garda, up to 1800 m), Holland (common), Norway (specimens
in coll. Strand, MP), and Iceland (MP). We could also re-examine the numerous
specimens from the Paris collection (many with labels bearing the name L. gallicus
Simon!). Figures based on specimens from Holland.

L. mengei thus seems to be absent from mediterranean Italy and from Spain,
Portugal and Greece. In the north it far exceeds the distribution of L. flavipes
(Blackwall), and the occurrence in Iceland, Lapland and Siberia shows an ability

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 11

to survive in subarctic regions. In the Alps (and Pyrenees?) it has been found at
higher elevations (up to 1800 m) than flavipes (up to 1200 m), demonstrating the
same climatic tolerance.

In Central Europe it is one of the commonest species of the tenuis group. It
occurs in leaf-litter, moss, sphagnum, and under stones. Adult specimens are
found at all seasons (Braun & Rabeler, 1969: 43; Palmgren, 1975: 60).

Lepthyphantes perseus Van Helsdingen, spec. nov.
(Fig. 6—8, 69)

Holotype: 4 , from Iran, Assalem, 2300 m, beech forest, v.1975, Ressl leg.; 23
paratypes from same locality (holotype in SMF, paratypes in personal collection
of J. Wunderlich and ML).

The three above specimens were entrusted to us for inclusion in our present
paper by Mr. J. Wunderlich of Neuenburg, Germany. Originating from a hardly
investigated region as Iran they are of great interest. The specimens much
resemble mengei, but differ in the shape of the lamella, terminal apophysis, and in
the dentition of the embolus. In size, coloration, cheliceral dentition, and
stridulating files there are hardly any differences between these species. On the
basis of the observed differences we treat this taxon as a separate species.

Measurements (in mm). Male. Total length 1.8—2.05, length cephalothorax
0.82—0.87. Legs: Fe I 1.20—1.35 times length cephalothorax.

A

7
8

Fig. 6—8. Lepthyphantes perseus Van Helsdingen, spec. nov. 6, male palp, lateral aspect; 7, radical
section, ventral aspect; 8, dentition on basal part of embolus, mesal aspect. 6,7, X 200; 8, X 740

12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

Coloration. Cephalothorax heavily suffused with grey, lateral margins and striae
darkest. Chelicerae with dark grey streaks on dorsal and lateral surfaces,
otherwise light brown. Sternum a shade darker, mouthparts with darker and
lighter areas, as chelicerae. Legs light brown to yellow-brown, not annulated;
palpal segments, the cymbium included, with strong, dark grey suffusion.
Abdomen with the usual faint male pattern of cross-bars of blackish grey colour,
intervening areas lighter grey with some white blotches (blotches absent in one
paratype); ventral and ventro-lateral surfaces dark grey.

Chelicerae with three dorsal teeth. Stridulating files fine, as in mengei (Fig. 69).

Chaetotaxy of legs not differing from the generic pattern. Position of d-spine
on tibia I 0.31—0.36, length of this spine 0.26—0.29 mm. Tm I 0.17—0.22.

Male palp (Fig. 6—8). Paracymbium with a single, rather obtuse, strong tooth
on the posterior margin of the proximal branch. Lamella with two equally long
branches, the dorsal one very gradually tapering to a slender tip, the ventral one
slightly less slender and shallowly forked at its tip (when viewed from dorsal or
ventral side); as in mengei, there is a ‘‘dorsal lobe” with a serrate margin at the
point where the element curves in anterior direction. Terminal apophysis with a
tongue-shaped projection which is shorter and relatively broader than in mengei.
Basal part of embolus with three or four denticles on a projecting mesal ridge.
Length of cymbium 0.27—0.29 mm.

The species is only known from the type-locality, where it was collected in a
beech forest, probably in leaf-litter.

Lepthyphantes flavipes (Blackwall)
(Fig. 9—12, 56)

Linyphia flavipes Blackwall, 1854: 178 (descr. 3 ; England).

Lepthyphantes flavipes; Miller, 1947: 40, Pl. 13 Fig. 2 (9). Locket & Millidge, 1953: 388, Fig. 231 B, D, F,
232 F (94 ). Wiehle, 1956: 205, Fig. 344—346 (93 ). Wanless, 1971: 23, 26, PI. 3, PI. 6 Fig. D (2);
1973: 129, 134, Pl. 1 Fig. 5, Pl. 6 Fig. 5, Fig. 24B (©).

Lepthyphantes tenebricola; Braun, 1960: 65 (misinterpretation of type-material of Linyphia tenebricola
Wider signalized but again misinterpreted); 1969: 216 (idem). Braun & Rabeler, 1969: 44 (idem).

Lepthyphantes zimmermanni; Fage, 1931: 181 (cave record from Rumania).

Lepthyphantes mengei; Denis, 1952: 12 (cave record from Rumania).

Theridium henricae Six, 1858: 294 (descr. 93 ; Netherlands).

Remarks. — To begin with the most disappointing discovery during our whole
revisionary work in this species-group, we have to admit here that we are not
convinced that figure 166 on plate 17 in Blackwall’s “A History of the Spiders of
Great Britain and Ireland” (part 2, 1864), where Linyphia flavipes Blackwall is
depicted, male and female, is the same as what is now unanimously called
Lepthyphantes flavipes (Blackwall). Especially the small figure of the male palp
positively can not belong to this species. The original description of Linyphia
flavipes by Blackwall (1854: 178) was not illustrated and the description does not
help much. Original material is not available. We are not the first to suggest this
discrepancy between the author’s original intention and subsequent interpretation.
Already Hull (1933: 106) pointed out that Blackwall’s species had been misinter-
preted. He also suggested (Hull & Harrison, 1937: 111) that Linyphia flavipes

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 13

Blackwall is the same as Bathyphantes pullatus (O. Pickard-Cambridge). He may be
right, but we prefer to leave the case as it is, because stability is served best, at
least in this case, by letting the matter rest. There is no end to Lepthyphantes
nomenclature if we start the game of replacing well established names by others.
Moreover, in most cases there is no solution to be offered with an acceptable
degree of certainty.

Braun (1960) published a remark on the discrepancy between the type-material
of Linyphia tenebricola Wider and Lepthyphantes tenebricola auct., though he can
not be held responsible for the re-identification of the type-series. The identifi-
cation of the original tenebricola specimens with Lepthyphantes flavipes (Blackwall),
as stated there, was also incorrect. See also under remarks on Lepthyphantes
tenebricola (Wider).

Fig. 9—12. Lepthyphantes flavipes (Blackwall). 9, male palp, lateral aspect; 10, radical section, ventral
aspect; 11, epigyne; 12, male chelicerae, anterior aspect. 9, 10, X 200; 11, X 190; 12, X 142

14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

The description of Theridium henricae by Six is very superficial and does not fit
any particular Dutch species. Subsequently, the name was used, by Chyzer &
Kulczynski (1894) for instance, for the species which in the British Isles was
known as Lepthyphantes flavipes and, later again, recognized as its synonym. The
question whether the name T. henricae was correctly applied to this species, is
difficult to answer. In the Rijksmuseum van Natuurlijke Historie at Leiden there
still exists a sample under this name in the Van Hasselt Collection. Van Hasselt
(1885: 180) stated that he had received material of T. henricae from Six (type-
material?), and that it was the same as Lepthyphantes tenebricola. (All other
specimens, however, identified by him with L. tenebricola belong either to
Lepthyphantes tenuis or to L. zimmermanni!). The sample of T. henricae consists of
juveniles of a Lepthyphantes species, which indeed could be L. flavipes, but it is
impossible to recognize the species with certainty. Besides, Chyzer & Kulczynski
(1894: 69—70) maintain, that they examined two male specimens of “Theridion
Henricae”, identified by Six and kept in the collection of Thorell. From their
remarks it becomes clear that this short series contained one specimen of mengei
and one of flavipes.

Diagnosis. — L. flavipes is among the smallest species of the tenuis group. It is
easily recognized by its coarse stridulating file (9), the shape of the epigyne, and
the cheliceral boss (4 ). Most specimens are darkly pigmented, especially the
chelicerae and the 5 palp.

Measurements (in mm). Total length, @ 1.7—2.6, 4 1.7—2.2; length cephalo-
thorax, 9 0.75—1.0, 3 0.77—0.90. Legs: Fe I 1.15—1.25 times length cephalo-
thorax in 9, 1.20—1.35 ing .

Stridulating files (Fig. 56) difficult to distinguish on the dark chelicerae (suffused
with grey); widely separated ridges present in female, absent in male, or very
indistinct and as widely separated as in female.!) Chelicerae of male (anterior
aspect) with a latero-basal boss and concave outer surface (Fig. 12).

Male palp (Fig. 9, 10) characterized by the single, relatively sharp (cf. mengei)
tooth on the posterior margin of the paracymbium, the shape of the lamella (with
the terminal branches standing about perpendicular on the subterminal part), and
the absence of denticles on the basal part of the embolus. The tooth on the
paracymbium is situated slightly more to the dorsal side than in mengei. The
terminal part of the embolus has a characteristic lobe-like extension on the latero-
ventral side (lateral aspect). The basal segments of the palp are strongly suffused
with grey, as are the chelicerae. Length of cymbium 0.27—0.30 mm.

Epigyne (Fig. 11) easily distinguished from that of the other species of this group
by the characteristically rounded basal part of the scape. Posterior median plate
hardly visible in the ventral aspect. There are two lateral depressions on the basal
part of the scape, but these are often very slight and hardly visible. Width of scape
0.140—0.175 mm.

1) This has already been pointed out in an earlier paper (Van Helsdingen, 1963: 36, Fig. 5, 6), but by
mistake the figure of the male chelicerae (Fig. 6) was said to be the female, and vice-versa.

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 15

Distribution. — According to Bonnet, the species is restricted to Central
Europe, including the British Isles; up to 1940 it was stated not to have been
recorded from Scandinavia north of Denmark (see, however, remarks below),
from Poland, the southern Balkans, or from Portugal. Since 1940 it has been found
in southern Finland, Poland, the European U.S.S.R., and Italy (cave).

It has been mentioned from caves in Hungary (Boker, 1922; Kolosvary, 1928)
Switzerland (Dresco, 1960, entrance), Italy (Brignoli, 1971), Spain (Fage, 1931),
and also occurs in caves in Rumania (Fage, 1931: 181, sub zimmermanni; Denis,
1952: 12, sub mengei; both re-examined, MP). See also Wolf (1934—1937).

We have examined material from Austria (North Tirol, common), Czechoslova-
kia (Böhmen, common), Yugoslavia (Slowenia, Hercegovina), Rumania (material
recorded by Niculescu, 1968), Italy (southern Alps, Parma, and Piemonte), Spain
(Prov. Huesca, Rodellar, leg. E. Duffey), and Holland (common). The figures are
based on specimens from Holland.

The following material has been re-examined: cave records from Spain by Fage
(1931) from the provinces Tarragona and Soria (MP); cave records from Rumania
by Denis (1952: 12, sub mengei; MP) and by Fage (1931: 181, sub zimmermanni;
MP); identified material in the Muséum National d’Histoire Naturelle, Paris.

Through the kindness of Dr. T. Kronestedt of the Naturhistoriska Riksmuseet,
Stockholm, a number of specimens could be examined, which are labelled
“Linyphia pygmaea Sundevall” and are part of the Westring collection. Among the
seven specimens examined, four were found to belong to flavipes. Westring’s
specimens are assumed to originate from Sweden, probably from the surroundings
of Göteborg, and thus we may have found an early proof of the existence of
L. flavipes in southern Sweden. It should be pointed out here that Chyzer &
Kulczynski (1894: 70) were already aware of the existence of L. flavipes (‘‘L. Hen-
ricae’’) in Sweden. They reported to have seen specimens of this species in the
collection of Thorell, partly under the name of Linyphia pygmaea (!). Bonnet
appears to have overlooked this important remark.

Thus L. flavipes appears to be indeed a Central European species, reaching its
northern limit in Great-Britain, Holland, Denmark, southern Sweden, southern
Finland, Poland and the European U.S.S.R.!) The upper limit in the Alps lies at
about 1200 m; in the Apennines there is a record from 1650 m (Parma). The
southern slopes of the Pyrenees, the Apennines, Slowenia, and Bulgaria form the
southern limit of its distribution, apart from the Spanish cave records from
Tarragona and Soria still more to the south.

It is a typical leaf-litter inhabiting species, in deciduous as well as coniferous
forests. Adults throughout the whole year (Braun & Rabeler, 1969: 44, sub
tenebricola).

1) There is a cryptic remark by Palmgren (1972: 91), who states to have found specimens of
Lepthyphantes zebrinus (Menge) in southern Finland, suggesting at the same time that this probably is a
geographical race (subspecies) of Lepthyphantes flavipes. This is the more surprising since Moritz (1968)
redescribed L. zebrinus and demonstrated a number of apparent differences between the two species.
In zebrinus the stridulating files are much finer, the epigyne has a medially excised scape, the
paracymbium of the male palp is toothless and of quite different shape, and the lamella does not show
any resemblance with that of flavipes. In our opinion zebrinus belongs to a different species-group.

16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

Lepthyphantes herbicola Simon
(Fig. 13-15, 57)

Lepthyphantes herbicola Simon, 1884: 323, Fig. 92 (descr. 9 3 ; France: Var, Alpes-Maritimes, Corse);
1929: 590, 595, Fig. 900-901 (3 9, diagnosis), 733 (references). De Dalmas, 1922: 88 (France: Ile
Giglio; cat.). Denis, 1933a: 572 (Pyrénées-Orientales); 1933b: 95 (Var); 1934: 152 (Var); 1935: 121
(Var); 1949: 18 (Provence).

Remarks. — It is not clear whether the original material of Simon is still extant.
The species was described in 1884 from the Departements Var, Alpes-Maritimes
and Corse. In the Paris Museum the available material from the time of Simon is
labelled Marseille and Banyuls, while there is also one series without a locality
label. A lectotype thus cannot be selected. There is no need for a neotype because
the species is not easily confounded with any of its congeners.

Fig. 13—15. Lepthyphantes herbicola Simon. 13, male palp, lateral aspect; 14, radical section, ventral
aspect; 15, epigyne. 13, 15, X 200; 14, X 180

Diagnosis. — L. herbicola belongs to the smaller species of the group and is well
characterized by its genitalia.
Measurements (mm). Total length, 9 1.8-2.3, 4 1.7-2.1; length cephalothorax,

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 17

© 0.77-0.92, 3 0.76-0.92. Legs: Fe I 1.1-1.3 times length cephalothorax in females,
1.15-1.30 times in males.

Stridulating files rather fine (Fig. 57), slightly coarser than in mengei.

Male palp (Fig. 13, 14) with only one small tooth on the paracymbium at the
anterior margin of the proximal branch (in contradistinction to mengei and
flavipes); the distal branch is much less slender than in the other species of the
group and appears to be rather strongly pigmented. The lamella is only slightly
curved and shows the usual dorsal and ventral arms. Embolus without denticles on
basal part. Length of cymbium 0.29-0.33 mm.

Epigyne (Fig. 15). Scape with gradually widening basal part with concave
margins; inner scape equally wide as basal part and also with concave margins;
posterior median plate distinct in the ventral aspect. As the specimens themselves,
the epigyne is small. Width of scape 0.125-0.150, of inner scape 0.125-0.140, of
posterior median plate 0.160-0.190 mm.

Distribution. — Literature records all refer to southern France and Corse; on
the mainland the species is restricted to the following provinces: Pyrénées-
Orientales, Bouches-du-Rhöne, Vaucluse, Var, Alpes-Maritimes, and Drome. In
1934, Denis stated it to be not uncommon (“Assez fréquent, sous les pierres et
dans les mousses’’) and Simon, too, indicates it to be common near Menton (1929).
Other samples come from moss and litter in mixed or coniferous forest.

We have studied all available identified material in the collection of the Paris
Museum (from Marseille, Banyuls, and an unlabelled series). New records can be
given here from:

France: Ardéche, Planzolles W. of Joyeuse, 700 m, open pine forest, 18.vii.1974,
P. J. van Helsdingen, 19 13; W. of Joyeuse, mixed pine-chestnut forest,
7.vii.1974, P. J. van Helsdingen, 19 (both ML).

Italy: Naples, 19 23, MP; Portici near Naples, 13, MP; Portici, 69, MP;
Isola Ischia near Naples, 19 15 , Mus. Bergamo.

Algeria: Edough, 59 lg , MP.

Spain: Prov. Huesca, Rodellar, 8.vi.1973, sifted from moss in gorge near Rio
Mascun, leg. E. Duffey and in his collection, 49 22 .

Yugoslavia: Dalmatia, Spilja Vranjaca, Kotlenice n. Split, S.vii.1971, 13 ; Isl.
Cres, Lipica jama, 1 9; both leg. C. Deeleman, ML.

All figures were made after specimens from Marseille.

Lepthyphantes tenuis (Blackwail)
(Fig. 16-18, 58)

Linyphia tenuis Blackwall, 1852: 18 (nom. nov. pro Linyphia pusilla Blackwall, preoccupied by Linyphia
pusilla Sundevall [= Microlinyphia pusilla (Sundevall)]).

Lepthyphantes tenuis; Locket & Millidge, 1953: 385, Fig. 230 E, 232 A (9 4 ). Wiehle, 1956: 197, Fig.
331-335 (9 4 ). Wanless, 1971: 22, 26, PI. 2 Fig. A, B, PI. 6 Fig. A (9); 1973: 129, 132, 134, Pl. 1 Fig.
1, Pl. 4 Fig. 7, Pl. 6 Fig. 1 (2).

Lepthyphantes falteronensis Di Caporiacco, 1936: 350 (descr. 9 2 : Italy, Apennines). Syn. nov.

Lepthyphantes aspromontis Di Caporiacco, 1949: 133, Fig. p. 134 (descr. 2 ; Italy). Alberti, 1967: 26 (type
catal. Mus. Triest). Syn. nov.

18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

Lepthyphantes sanfilippoi Di Caporiacco, 1950: 108, Fig. 2 (déscr. 3 ; Italy, Liguria, in cave). Syn. nov.
?Linyphia arctica Keyserling, 1886: 85, Pl. 14 Fig. 179 (descr. © ; Sitka, Alaska).
Lepthyphantes zimmermanni; Fage, 1931: 181 (Portugal, cave).

Remarks. — The name tenuis was introduced by Blackwall as a replacement
name for his pusilla in the genus Linyphia, which name had already been used in
this genus as a primary combination by Sundevall. We have not located any
original material of Blackwall’s Linyphia pusilla.

The original material of Lepthyphantes falteronensis Di Caporiacco could be
located in the collection of Di Caporiacco in the Museo Zoologico in Firenze
(MZF). It consists of 2 4 from Falterona and 2 3 from Monte Acuto. Not all
specimens are in good condition: from either locality one specimen lacks both
palps, the second specimen from Monte Acuto has a left palp only, the material
from Monte Falterona is accompanied by a loose palp. The only complete
specimen (from Monte Falterona) is designated lectotype here. The type-locality,
Monte Falterona, is situated ENE of Florence in the Apennines. The specimens
were collected at 1650 m (Mt. Falterona) and 1428 m (Mt. Acuto), respectively.

The male holotype of Lepthyphantes aspromontis Di Caporiacco, described from
Italy (Calabria, Aspromonte), could be examined through courtesy of Dr. G.
Alberti of the Museo Civico di Storia Naturale at Trieste, where the type is
preserved. It belongs to L. tenuis (Blackwall).

Of the original material of Lepthyphantes sanfilippoi Di Caporiacco only one
specimen could be located. It is selected as the (male) lectotype here. It is
preserved at the Museo Civico di Storia Naturale at Milano and could be
examined through the kindness of Dr. C. Leonardi. It appears to belong to
Lepthyphantes tenuis (Blackwall).

The original material of Linyphia arctica Keyserling (1 @ only?) could not be
located at the AMNH, nor at the BM; it is probably lost. Our suggestion of a
possible synonymy is mainly based on the small figure of the epigyne while
measurements and other details given by Keyserling agree with tenuis.

Specimens from la Palma, Canary Islands, are said to be different from the
normal form in the shape of the lamella, the dentition of the paracymbium, and
the coloration (Schmidt, 1975b). We have not seen the specimens.

Diagnosis. — A comparatively light species of small size. The paracymbium has
a very characteristic pair of denticles, the lamella is easily recognized by its shape,
the scape of the epigyne has the shape of an anchor.

Measurements (in mm). Total length, 9 2.3-2.8, 3 1.95-2.4; length cephalo-
thorax, 9 0.95-1.10, + 0.82-1.02. Legs: Fe I 1.25-1.35 times length cephalothorax in
females, 1.25-1.4 times in males.

Stridulating files rather coarse (Fig. 58).

Male palp (Fig. 17, 18) easily recognized by the dentition of the paracymbium
and the rather blunt dorsal branch of the lamella. Paracymbium with a small and
sharp denticle at the posterior margin and a second, barely larger and slightly
more conical denticle at the anterior margin of the proximal branch; posterior
denticle pointing outward and backward, more outward than in zimmermanni.
Lamella relatively small, the dorsal branch ending well before ventral branch, with

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 19

Fig. 16—18. Lepthyphantes tenuis (Blackwall). 16, epigyne; 17, radical section, ventral aspect; 18, male
palp, lateral aspect. 16, X 194; 17, 18, X 180

obtuse tip; ventral branch slender and rather straight, sometimes forked at tip.
Embolus with well-developed cluster of small teeth on basal part, numbering at
least 7. Length of cymbium 0.29-0.34 mm.

Epigyne (Fig. 16) with a distinctly anchor-shaped scape; anterior section more
or less parallel-sided or with concave margins, gradually diverging in posterior
direction but not fluently passing into margins of broadest part of visible scape;
inner scape with conspicuously rounded lateral lobes, about as wide as broadest
point of exposed scape; posterior median plate hardly wider than scape. Width of
scape 0.135-0.165, of inner scape 0.125-0.150, of posterior median plate 0.160-0.190
mm.

1) It has been suggested by Simon (1929: 733, footnote 2) that Linyphia (Lepthyphantes) taczanowskii O.
P.-Cambridge, 1873 [misprinted tarzanowskii] would be close to tenuis. However, re-examination of the
original material, which is still preserved in the collections of the Hope Department of Zoology
(Entomology) at Oxford, revealed it to be a senior synonym [syn. nov.] of Lepthyphantes trucidans (L.
Koch, 1879), described from the Yenissey region in Siberia. Linyphia taczanowskii was also collected in
Siberia (Lake Baikal). L. trucidans does not fit in with the tenuis group.

20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

Distribution. — For Lepthyphantes tenuis, Bonnet listed references for nearly all
European countries with the exception of Iceland, Norway, Sweden and Finland.
Russia and the Caucasus!) formed the eastern limit, Greece, Italy, Spain and
Portugal the southern limit on the European mainland. Several records were
available from the Acores, Madeira and New Zealand, but none from mediter-
ranean North Africa.

Since 1940, this distribution was confirmed for many countries, while new
countries could be added to it: Afghanistan (Denis, 1958) and Finland (Prov.
Vaasa, 63°N, Hackman, 1951; Tvärminne, southern Finland, Palmgren, 1972,
1975). The occurrence of this species in the Canary Islands was established by
Schmidt (1975 a, b), the occurrence on Madeira and the Acores was confirmed by
Denis (1962 a and 1963, and 1964, respectively). Recently (Bragg & Leech, 1972) it
has been recorded also from the Nearctic region (British Columbia, © only), from
where it probably had been recorded already by Keyserling (1886).

The species has been found in caves in Belgium (Leruth, 1935, 1939), France
(Denis, 1959: Pyrenees), Portugal (Fage, 1931: sub zimmermanni, re-examined),
Italy (Brignoli, 1971), and Bulgaria (Drensky, 1931; Deltshev, 1972 a, 1972 b, 1973).
See also Wolf (1934-1937).

Samples have been re-examined from the Acores (Berland, 1932), New Zealand
(De Dalmas, 1917), and Portugal (Fage, 1931: 181, zimmermanni 13 , belongs to
tenuis) (all MP).

Material has been found from the following countries, so that the known range
can be extended to: Algeria (several series from Edough, Constantine and Setif;
MP), Lebanon (MP), Norway (Bergen; MP).

The diagnosis, measurements and figures are based on material from Holland
(common), France (Menton), Czechoslovakia (not uncommon, lowland), Bulgaria
(cave), and Italy (type-series of L. falteronensis and L. aspromontis). Specimens
from Greece (Salonique, MP) and Spain (Prov. Huesca, Rodellar; BM) were also
used.

With records from Madeira, the Canary Islands, Algeria, the Lebanon, and
Afghanistan, L. tenuis reaches further south and south-east into warm, subtropical
regions than most members of the tenuis group. It does not occur far to the north
and appears to be rare there. In the Alps it is restricted to lower altitudes.

In New Zealand it must have been introduced. At least this is the only
reasonable explanation for the extremely wide gap in the distribution. According
to R. R. Forster (in litt.) it is extremely common in open country and in and
around gardens in settled areas, but it does not occur in native forests. It also
seems to be spreading to other islands in the south. For the Nearctic region an
introduction is less self-evident. Recently 3 © have been mentioned from W.
Canada (Bragg & Leech, 1972, British Columbia), while we (vH) also received for
examination a © specimen from the State of Washington, through the kindness of
Dr. Dondale, Ottawa. If Linyphia arctica Keyserling is indeed based on a specimen
of tenuis, as suggested above, the species has been found also in southern Alaska
(Sitka). Importation of spiders by man in the Nearctic Region is reported on by
Lindroth (1957), on the east as well as on the west coast. However, we should be
very careful with hasty conclusions.

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 21

L. tenuis does occur in the litter stratum in forests, but it is equally common in a
wide variety of other habitats. In Holland it is, among other habitats, very
common in tussocks of Marram grass on the seadunes. Adults throughout most of
the year (Braun & Rabeler, 1969: 44).

Lepthyphantes tenebricoloides Schenkel
(Fig. 19-21, 59)

Lepthyphantes tenebricoloides Schenkel, 1938: 15, Fig. 5 a-e (descr. 9 & ; Madeira). Denis, 1941: 113
(Tenerife); 1962 a: 77 (Madeira). Forcart, 1961: 72 (catal.). Schmidt, 1975 a: 224 (Canary Islands;
aberrant specimens?); 1975 b: 235 (Canary Islands).

Remarks. — The original description by Schenkel is lengthy and detailed and
includes remarks on the variation in abdominal patterns. The original material
should be at the Naturhistoriska Riksmuseet at Stockholm. We have only seen 39

and 33 paratypes from the Schenkel collection at Basel (not 29 and 43 as stated
by Forcart, 1961: 72).

Fig. 19—21. Lepthyphantes tenebricoloides Schenkel. 19, epigyne; 20, male palp, lateral aspect; 21,
radical section, ventral aspect. 19, X 190; 20, 21, X 160

22 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

According io Schmidt (1975 a), recently collected material from Gomera,
Canary Islands, differs considerably from the description by Schenkel, especially
as to the paracymbium. In the Madeira specimens the paracymbium is toothless;
those from Gomera are said to show some dentition, but the position of the teeth
is not indicated.

Diagnosis. — L. tenebricoloides is very light: the cephalothorax a very light
yellow-brown with a narrow median dark grey streak that widens to the breadth of
the eye-region in front; the abdomen gives a light impression and bears pairs of
black spots dorsally, though the darker parts can be more pronounced in some
specimens (see Schenkel’s description and figures). On the ventral surface of the
abdomen there are a pair of latero-ventral light spots halfway between the
epigastric furrow and the spinnerets.

Measurements (in mm). Total length, 9 2.6-4.1, 3 2.6-3.0; length cephalo-
thorax, 9 1.2-1.55 (Schenkel mentioned a female cephalothorax length of 1.7), 3
1.2-1.37. Legs: Fe I 1.1-1.2 times length cephalothorax in females, 1.2-1.35 times in
males.

Stridulating files (Fig. 59) rather fine, conspicuous and well-developed in the
males, less conspicuous in the females. Dorsal margin of chelicerae with three
teeth as usual, but teeth long and slender, distinctly longer than in other species of
this group.

Male palp (Fig. 20, 21). Paracymbium without any teeth. Lamella with a nearly
straight ventral margin, the ventral branch projecting straight forward and slightly
dorsal, the dorsal branch rounded-truncated. Base of embolus without denticles.
Length of cymbium 0.35-0.39 mm.

Epigyne (Fig. 19) with basal part of scape gradually widening with concave
margins towards broadest point; lateral lobes of inner scape as wide as basal part,
their lateral margins about straight and converging in anterior direction; posterior
median plate slightly wider than scape. Width of basal part of scape 0.150-0.200, of
inner scape 0.160-0.175, of posterior median plate 0.185-0.225 mm.

Distribution. — L. tenebricoloides was described from Madeira. It has also been
recorded from the Canary Islands (Denis, 1941). There are no records outside this
archipelago. Recently again recorded from the Canary Islands (Schmidt, 1975 a,
b), but the specimens are reported to disagree with the original description.

Lepthyphantes drenskyi Van Helsdingen, spec. nov.
(Fig. 22, 23, 60)

One sample of this new species was found in what is left of Drensky’s collection
in the Zoologisches Institut der Bulgarischen Akademie der Wissenschaften,
Sofia; it had been identified (by Drensky?) with L. tenebricola. However, it appears
to be different from tenebricola and rather reminds of tenuis, from wich it differs in
the dentition of the paracymbium, the number of denticles on the embolus, and
the stridulating files. There are no adult females in the sample.

Types. — 4 holotype from Dragalewzi near Sofia, vi.1909, P. Drensky.

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 23

22 23

Fig. 22—23. Lepthyphantes drenskyi Van Helsdingen, spec. nov. 22, male palp, lateral aspect; 23, radical
section, ventral aspect. X 142

Paratypes (13 , 1 subadult 3 , 1 subadult 9) from the same locality, preserved with
the holotype. All specimens are in the Zoologisches Institut at Sofia.

Description. — Measurements (in mm). Male. Total length 2.2-2.4, length
cephalothorax 0.95-1.05. Fe I 1.25-1.40 times length cephalothorax.

Coloration. The specimens are light yellow-brown (discoloured through pre-
servation?). Dorsal pattern distinct, composed of greyish chevrons.

Chelicerae with coarse stridulating files (Fig. 60). Legs with the usual spines,
position of the retrodorsal (basal) spine on tibia I 0.30-0.31, length of this spine
0.325 mm. Tm 10.22. palp (Fig. 22, 23). Paracymbium with a single, small denticle at
posterior margin. Embolus with four denticles on the protruding part of the basal
half. Lamella not unlike that of tenuis, but the dorsal branch more truncate, blunt,
not pointed. Length of cymbium 0.38-0.40 mm.

As indicated above, L. drenskyi is not unlike tenuis, but in that species the
paracymbium bears a second tooth at the anterior margin of the proximal branch,
the embolus has seven or more teeth, and the dorsal branch of the lamella is short,
but distinctly pointed, not truncate. The stridulating files make a coarser
appearance.

The species is named after Pentcho Drensky, as a late homage to this well-
known Bulgarian arachnologist, who has contributed so much to our knowledge of
the Bulgarian spider fauna, and who also collected the type-series.

Lepthyphantes zimmermanni Bertkau
(Fig. 24-27, 61)

Lepthyphantes zimmermanni Bertkau, 1890: 10 (nom. nov. pro Lepthyphantes zebrinus Simon, 1884).
Locket & Millidge, 1953: 386, Fig. 230 B, C, 231 G, 232 B (9 g ). Wiehle, 1956: 200, Fig. 336-339 (©
& ). Wanless, 1971: 22, 26, PI. 2, PI. 6 Fig. B (9); 1973: 129, 134, PI. 1 Fig. 4, Pl. 6 Fig. 4 (9).

24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

Lepthyphantes borealis Braendegärd, 1932: 12, Fig. 2 (descr. 9; Iceland); 1958: 84, 85 (= variety of zim-
mermanni).
Lepthyphantes cristatus; Simon, 1892: clxxvii (Iceland).

Remarks. — Simon (1929: 595) described a variety spiniger from Southern
France, which subsequently has been recognized as a separate species (Dresco &
Jézéquel, 1961). It is treated as such in this paper.

Diagnosis. — The male of this species is well characterized, apart from the
shape of the lamella, by the peculiar small and sharp backward pointing denticle

Fig. 24—27. Lepthyphantes zimmermanni Bertkau. 24, epigyne (specimen from Holland); 25, epigyne

(specimen from Bulgaria); 26, radical section, ventral aspect; 27, male palp, lateral aspect. 24, X 165;
26, 27, X 160; 25, X 142

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 25

on the posterior margin of the paracymbium. The epigyne is much like that of
L. floriana, but there are differences in the shape of the scape and the laterally
protruding wings of the underlying arched part of the latter species.

Measurements (in mm). Total length, 9 2.2-3.1, 4 2.2-2.75; length cephalo-
thorax, @ 0.95-1.25, 3 1.0-1.25. Legs: Fe I 1.2-1.4 times length cephalothorax in
females, 1.15-1.35 times in males.

Stridulating files (Fig. 61) coarse at base to rather fine toward apex, well-
developed, slightly coarser than in floriana from Rumania.

Male palp (fig. 26, 27) characterized by the single small but sharp tooth near the
posterior margin of the paracymbium, which points backward rather than
outward. Lamella evenly curved, with a distinct separate dorsal branch and a
much longer ventral branch, the latter slender and tapering to a sharp tip. Base of
embolus on ventral side with seven denticles (cf. spiniger). Length of cymbium
0.37-0.43 mm.

Epigyne (Fig. 24) with a long and slender anterior section of basal part of scape,
parallel-sided, rather suddenly widening into the much broader posterior section;
lateral wings of inner scape rather angular, margins in front of projecting angle
rather straight; inner scape as wide as basal part; posterior median plate distinctly
wider than scape (shape of median plate much depending on angle of vision).
Width of basal part of scape 0.150-0.175, of inner scape 0.150-0.175, of posterior
median plate in ventral view 0.210-0.250 mm.

Morphological variation. A slightly aberrant specimen from Bulgaria (9, Fig.
25) is tentatively placed here, though the ventral aspect of the scape is obviously
different; also the posterior aspect reveals a swollen ventral surface of the basal
part of the scape, while this part is much flatter in zimmermanni. It is unfortunate
that we do not have any zimmermanni male from Bulgaria, or a good sample of this
type of female with matching males.

Distribution. — The records up to 1940, as given by Bonnet, cover most of
Europe, including the British Isles but with the exception of Norway, Sweden and
Finland, Poland and more to the east. There are, however, records from the
Faroers, and also from Iceland, where it seems well-established (Braendegärd,
1958: 85). In Southern Europe there are records from Portugal (Fage, 1931: 181: =
tenuis, re-examined), from Spain (Galiano, 1910, sub zebrinus, citing a record by
Simon; Denis, 1938; Fage, 1931, cave record, re-examined), and from Italy
(several authors). The Balkans and Hungary formed the eastern boundary in 1940.
The original description of Bathyphantes pygmaeus Menge is considered by some
authors to refer, at least partly, to zimmermanni; in that case Poland (Danzig) falls
within the distribution of the species as known up to 1940.

Since 1940, there are new records, among others, from Poland, European
U.S.S.R., Bulgaria, Finland and Ireland. There is one record from northern
Yugoslavia (Polenec, 1958).

Cave records come from Belgium (Leruth, 1935, 1939), France (Fage, 1931, re-
examined; Denis, 1959, Pyrenees; Denis, 1967, Haute-Garonne), from Spain
(Fage, 1931, re-examined), Bulgaria (Deltshev, 1972 a, 1972 b, and 1973; specimen
re-examined and identification maintained, despite observed differences (Fig. 25)).

26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

The cave records from Portugal (Fage, 1931: 181, = tenuis, re-examined) were
based on misidentifications. See also Wolf (1934-1937).

We have examined a fair number of specimens from Switzerland (coll.
Schenkel) and France (coll. Simon). A specimen in the collection of Simon,
originating from Iceland and identified with L. cristatus (Menge) (Simon, 1892 2),
was found to belong to zimmermanni. Other examined material came from
Czechoslovakia (one locality only), Bulgaria (1 © only, in cave), and Holland. In
Tirol and eastern Switzerland it seems to be lacking. The distribution in the
Balkans is hardly known. We presume zimmermanni to be absent from higher
mountainous regions. Our figures are based on specimens from Holland.

L. zimmermanni is found in the litter stratum of preferably coniferous forest
(Braun & Rabeler, 1969: 44).

Fig. 28—30. Lepthyphantes spiniger Simon. 28, male palp, lateral aspect; 29, radical section, ventral
aspect; 30, epigyne. 28, 29, X 160; 30, X 200

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 27

Lepthyphantes spiniger Simon
(Fig. 28-30, 62)

Lepthyphantes zimmermanni spiniger Simon, 1929: 595 (descr.; Pyrenees and Alps), 733 (reference).
Fage, 1931: 181, 237 (France: cave in Ariège). Dresco, 1949: 187 (France: Pyrénées-Centrales).
Denis, 1957: 254 (Pyrénées).

Lepthyphantes spiniger; Dresco & Jézéquel, 1961: 105, Fig. 1-4 (deser. 3 9, distinct species; Basses-
Pyrénées, Hautes-Pyrénées, Haute-Garonne).

Remarks. — The collection of the Museum National d'Histoire Naturelle at
Paris contains a number of samples of this species, but it is not clear which of these
belongs to the material on which Simon based his diagnostic remarks. All samples
contained specimens of other species as well. Still we do not think it necessary to
select a lectotype, because the species is quite characteristic and can hardly be
confounded with any of the group members.

Diagnosis. — L. spiniger is most easily distinguished from its congeners by the
spinose metatarsi and the very fine stridulating files of the chelicerae.

Measurements (mm). Total length, 9 2.7-3.2, & 2.1-2.7; length cephalothorax,
Q 1.10-1.27, ga 1.10-1.20. Legs: Fe I 1.15-1.3 times length cephalothorax in
females, 1.25-1.45 times in males.

Stridulating files (Fig. 62) very fine in both sexes. Metatarsi with a l’- and l”-
spine beside the usual dorsal spine, but metatarsus III not always bearing a
complete set.

Male palp (Fig. 28, 29) characterized by the large lamella, which has the longer
terminal branch forked, and by the large number of denticles (11) on the basal part
of the embolus. The paracymbium bears a single small tooth, which does not point
backward as in zimmermanni but dorsad, while below it a ridge-like excrescence is
found along the posterior margin. Length of cymbium 0.36-0.41 mm.

Epigyne (Fig. 30) resembling that of zimmermanni, but posterior section of basal
part of scape more clearly showing the reversal of its course, the lateral margins
distinctly curving anterad; lateral wings of inner scape wide and broadly rounded;
entire scape slightly more protruding posterad. Posterior median plate less visible
than in zimmermanni. Width of basal part of scape 0.150-0.190, of inner scape
(measured between apices of wings) 0.160-0.175, of posterior median plate 0.185-
0.225 mm.

Distribution. — The species, according to Simon, 1929 (p. 733), occurs in
coniferous forests in the Pyrenees and the Alps. Actually this can only be
confirmed for the Pyrenees. We have examined all available material at the
Museum National d’Histoire Naturelle at Paris, which came from Bagnères-de-
Bigorre (Hautes-Pyrénées), from the ‘Basses-Pyrénées”, from the Grotte
d’Aubert (Ariege: Moulis), and from the Grotte de Lestelas (Ariege: St. Lizier)
(Fage, 1931). Apart from Simon’s reference to the Alps, all other authors gave
records from the Pyrenees only. Of four tubes in the Paris Museum the labels bear
only numbers, which appear to be of no help in establishing the origin of the
material. These samples might indeed come from the Alps, but we should like to
have confirmed the occurrence in this region by fresh and well-localized captures.

28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

Nearly all samples referred to above were a mixture of spiniger and zimmerman-
ni, in several cases mixed up with other Lepthyphantes species as well.

The species was collected in the entrances of caves and in dolines, but also
outside caves in the litter stratum of coniferous forests.

Lepthyphantes floriana Van Helsdingen, spec. nov.
(Fig. 31-34, 63)

Lepthyphantes zimmermanni; Niculescu-Burlacu, 1968: 91 (Rumania).

Types. — Holotype g from Branesti, near Bucarest, Rumania (ML); many 9
and g paratypes from the same locality (ML and Institutul de Biologie “Traian
Savulescu” at Bucarest).

Specimens of this species were doubtfully referred to L. zimmermanni by one of
us (vH) when he advised Miss Burlacu in 1967 during her faunistic research in the
Branesti wood near Bucarest. In the course of our present review we realized that
a set of constant differential characters neatly separates this population from
zimmermanni and the other species of this group, and that we are dealing with a
separate species of the zimmermanni subgroup. It is a pleasure to name this species
after Mrs. Floriana Niculescu-Burlacu.

Diagnosis. — A species of the tenuis group, which closely resembles zimmerman-
ni but differs in the dentition of the paracymbium, having two teeth instead of one;
the shape of the lamella is also different. The lateral wings of the inner scape in the
epigyne are of more angular shape than in zimmermanni and spiniger. The
stridulating files (Fig. 63) are finer, with the ridges closer together than in
zimmermanni, but not as fine as in spiniger. The legs, especially the femora, are
conspicuously whitish yellow.

Measurements (mm). Total length, 9 2.5-2.7, 2.0-2.6; length cephalothorax,
Q & 1.0-1.1. Legs: Fe I 1.15-1.3 times length cephalothorax in females, 1.2-1.3
times in males.

Male palp (Fig. 31-33) with the paracymbium furnished with two teeth at the
posterior margin, the upper one pointing downward, the lower one slightly larger
and pointing upward, thus enclosing a roundish dent between them. Lamella with
a dorsal lobe above the main lateral branch, dorsal and ventral branches after the
curve rather short. Base of embolus with few denticles or none, this character
being variable. Length of cymbium 0.40-0.44 mm.

Epigyne (Fig. 34) hardly differing from that of zimmermanni, but the wing-like
extensions of the inner scape with a distinct posterior angle and often with an
additional projection just in front of broadest point, giving the wings an angular
appearance. Posterior median plate largely covered by the wings of the inner
scape and thus less conspicuous than in zimmermanni. Width of basal part of scape
0.175-0.190, of inner scape (measured between tips of wings) 0.185-0.200, of
posterior median plate 0.210-0.240 mm.

Distribution. — L. floriana is described after specimens from Branesti forest
near Bucarest, Rumania, where it appears to be very common. The specimens

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 29

were collected with Barber traps and inhabit the leaflitter stratum of deciduous
forest.

A single © specimen from Drensky’s collection (Zoological Institute, Sofia)
seems to belong to this species; it originates from Tscherepischki monastir near
Iskar, Bulgaria, vi.1916, leg. P. Drensky, which suggests a wider distribution in the
Balkans.

Fig. 31—34. Lepthyphantes floriana Van Helsdingen, spec. nov. 31, male palp, lateral aspect; 32, radical
section, ventral aspect; 33, paracymbium; 34, epigyne. 33, X 142; 34, X 150; 31, 32, X 160

30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

Lepthyphantes tenebricola (Wider)
(Fig. 35-38, 64)

Linyphia tenebricola Wider, 1834: 267, Pl. 18 Fig. 2 (descr. 9 ; Germany).

Lepthyphantes tenebricola; Miller, 1947: 40, PI. 13 Fig. 3 (9). Locket & Millidge, 1953: 388, Fig. 230 D,
232 G (9 2). Wiehle, 1956: 145, Fig. 327-330 (9 g ). Wanless, 1971: 23, 26, PI. 3, PI. 6 Fig. C (9);
1973: 129, 134, PI. 1 Fig. 2, PI. 6 Fig. 2 (9).

Linyphia arcuata Thorell, 1856: 168 (descr. 9 3 ; Sweden).

Lepthyphantes arcuatus; Braun, 1960: 65 (used as oldest available synonym because the type-material of
Linyphia tenebricola Wider was found to belong to another species; see, however, under remarks
below); 1969: 215 (idem). Braun & Rabeler, 1969: 41 (idem).

Remarks. — Braun (1960), having been informed that the type-series of Linyphia
tenebricola Wider (in SMF) did not belong to the species currently recognized
under that name (in the combination Lepthyphantes tenebricola), but to Lepthyphan-
tes flavipes (Blackwall), solved the problem by replacing flavipes Blackwall
(from 1854) by the older tenebricola Wider (from 1834). Lepthyphantes tenebricola
auct. consequently had to be replaced by the oldest available synonym, Linyphia

Fig. 35—38. Lepthyphantes tenebricola (Wider). 35, radical section, ventral aspect; 36, male palp, lateral
aspect; 37, epigyne (specimen from French Jura); 38, epigyne (specimen from Holland). 37, X 135; 35,
36, X 144; 38, X 158

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 31

arcuata Thorell, 1856. Subsequently it was found that the one misidentification had
been replaced by another (Locket, Millidge & Van Helsdingen, 1970: 90), as the
type-specimens were found to belong to Lepthyphantes mengei Kulczynski, 1887,
and not to L. flavipes (Blackwall). Straightforward nomenclatorial procedures
would ask for another shifting of names, replacing mengei Kulczynski (from 1887)
by tenebricola Wider (from 1834), using arcuatus Thorell (from 1856) for tenebricola
auct., and re-establishing flavipes Blackwall in its original sense. However, it is
thought highly unadvisable to carry through such a shifting of names for so
common and frequently used names — not only by taxonomists. A proposal to
stabilize the situation has therefore been presented to the International Commis-
sion on Zoological Nomenclature (see Locket, Millidge & Van Helsdingen, 1970).
We deal with the species in the current sense here.

The synonymy of Linyphia arcuata Thorell with the species currently referred to
as Lepthyphantes tenebricola (Wider) is difficult to ascertain, unambiguous type-
material of arcuata not being available.

Diagnosis. — The species is easily recognized by the dentition of the
paracymbium and the relatively simple lamella. The epigyne has a comparatively
broad basal portion of the visible scape.

Measurements (in mm). Total length, 9 2.2-2.8, 4 2.15-2.35; length cephalo-
thorax, 9 0.92-1.12, & 0.97-1.05. Legs: Fe I 1.1-1.2 times cephalothorax in
females, 1.1-1.25 times in males.

Stridulating files (Fig. 64) coarse at base, finer at apex.

Male palp (Fig. 35, 36): paracymbium with two teeth on the basal branch, viz., a
blunt one near the posterior margin and a long and slender one behind the distal
branch when seen from the side, but revealed in full length when viewed slightly
from behind. Lamella sickle-shaped, its outer (dorsal) curve with or without an
additional branch, of comparatively simple shape. Base of embolus with 3 to 7
denticles. Length of cymbium 0.36-0.39 mm.

The variation observed in the palp of tenebricola concerns the size of the large,
distal tooth on the paracymbium, the shape of the lamella, and the number of
denticles on the base of the embolus. As to the lamella, it has already been pointed
out above that the outer margin may have a separate short branch dorsally,
otherwise this branch is only indicated by a small denticle. The number of
denticles on the basal section of the embolus varies from three to seven.

Epigyne (Fig. 38) with basal part of scape gradually widening from the relatively
broad anterior section (two-fifths to one-half of maximum width of scape) to the
broadly rounded posterior margin where the scape turns dorsad and inward; at this
point the scape is conspicuously dark brown, a sign of heavy sclerotization,
distinguishing it at once from most other species recognized so far; lateral wings of
inner scape with tips as far apart as width of basal part of scape or slightly less, and
rounded, never angular. Posterior median plate visible at either side of scape, but
not very conspicuously so. Width of basal part of scape 0.175-0.215, of inner scape
0.150-0.190, of posterior median plate 0.200-0.240 mm.

The epigyne quite obviously is subject to a relatively large variation. Not only
the ratio width anterior section to width posterior section of the basal part of

32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

the scape varies between 0.4 and 0.5 (which makes quite a different impression),
but there also is a large amount of variation in the development of a median pit or
depression at half length of the basal part of the scape. This pit can be quite
conspicuous because of the stronger pigmentation (sclerotization) of the surroun-
ding area (rare), or it merely consists of a slight depression with only the slightest
pigmentation. We have seen well pitted scapes in specimens from the French Jura
(Fig. 37) and from Graubunden in Switzerland. The lateral wings of the inner
scape are rounded with a straight or slightly concave margin in front of the
broadest point.

Distribution. — Bonnet could list references for nearly the whole of Europe,
with the exception of Iceland and Greece. Moreover, the species had been found
on the Acores, Madeira, and in Kamtchatka.

Since 1940, the species has again been recorded from many countries in Europe.
The only record from Madeira (Simon, 1897), based on a single adult female (MP,
examined), was already disputed by Denis (1962: 77, 108) and is here referred to
tenuis. We have not seen the specimens from the Acores. The material from
Kamtchatka certainly needs to be re-examined.

The species is said to occur also in caves (records from Switzerland and
Hungary, see Wolf, 1934-1937). It is certainly not a troglophilous species and may
have been found near the entrances only.

Our observations are based on material from Austria (common), Czechoslova-
kia (common), Switzerland (common), Italy, Bulgaria and Holland (infrequent).
Figures based on specimens from Holland. It is a widespread species and usually
referred to as not uncommon in the leaf-litter stratum in forests. It is common in
the Swiss and Austrian Alps (up to 2,000 m). Most adult specimens were collected
in the summer (Braun & Rabeler, 1969: 41, sub arcuatus; Palmgren, 1975: 59).

Lepthyphantes jacksoni Schenkel
(Fig. 39-42, 65)

Lepthyphantes jacksoni Schenkel, 1925: 266, 302, Fig. 10 A-D (deser. 9 ¢ ; Wallis, Switzerland); 1927:
229 (Wallis); 1929: 10 (Tessin); 1933: 15 (Wallis). Forcart, 1961: 72 (designation of lectotype).
Lepthyphantes tenebricola; Schenkel, 1929: 10 (p.p.; Tessin).

Types. — The original material of this species is preserved in the Basel
collection, together with that of later records. A male, bearing the label “Weg vom
Hotel zum Märjelensee’ was selected lectotype by Forcart (1961). A female from
“Weg vom Hotel Jungfrau zum Eggishorn” is the only paralectotype.

Diagnosis. — L.jacksoni belongs to the species-subgroup with a pit or
depression in the scape of the epigyne and a single large tooth on the
paracymbium. It can be distinguished from the other member of the subgroup in
the Alps by the shape of the lamella which has the two branches clearly diverging;
in jacksonoides the branches run parallel.

Measurements (in mm). Total length, @ 2.2-2.5, g 1.95-2.2; length cephalo-
thorax, 9 0.96-1.08, 4 0.90-1.08. Legs: Fe I 1.15-1.25 times length cephalothorax in
females, 1.1-1.2 times in males.

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 33

Stridulating files rather coarse (Fig. 65).

Male palp (Fig. 39-41). Paracymbium with a single, fair-sized tooth close to the
inner margin of the distal branch and thus not visible in the lateral aspect, but
quite conspicuous in postero-lateral view. Lamella forked, both branches well-
developed, diverging (cf. jacksonoides), dorsal branch pointing forward, ventral
branch longest and curved antero-ventrally. Terminal apophysis with a well-
developed and sclerotized ventral lobe, which is blade-like and curved upward.
Base of embolus with 1-4 denticles. Length of cymbium 0.42-0.45 mm.

Fig. 39—42. Lepthyphantes jacksoni Schenkel. 39, male palp, lateral aspect; 40, paracymbium; 41,
radical section, ventral aspect; 42, epigyne. X 142

Epigyne (Fig. 42). Basal part of scape with parallel-sided stem, widening into the
much broader posterior section which bears a characteristic, oval depression just
in front of the transition of basal part into inner scape; lateral wings of inner scape
rounded or slightly angular, inner scape at this point nearly as wide as broadest,
posterior section of basal part. Posterior median plate distinctly wider than scape.
Width of basal part of scape 0.200-0.215, of inner scape 0.185-0.190, of posterior
median plate 0.250 mm. In the lateral aspect the strong curvature of the basal part
of the scape and the depression near the tip are distinct.

34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977 |

Distribution. — Up to the present the species had only been recorded by |
Schenkel from Wallis and Tessin in Switzerland. All Schenkel’s specimens (see |
below) were collected in July and August, and above 2,000 m, but the number of |
records is too scanty yet to consider these data as the absolute limits in time and |
distribution. It is not unlikely that, as in jacksonoides, the species lives outside
forests in the herbal stratum, up to 50 cm above the soil, rather than close to the
ground in leaf-litter, moss, and between the lower parts of grasses and herbs.

In the eastern Alps, L. jacksoni seems to be replaced by another species of this
subgroup, L. jacksonoides. The latter has been found at 1,500 m and above in
northern and eastern Switzerland, the Austrian Alps, and in Bulgaria. We do not |
know if and where the distributions of the two species meet or overlap. In fact, this |
kind of information might serve to evaluate the characters here used to distinguish
between the two species. |

All specimens mentioned by Schenkel in subsequent years have been re-
examined. The only additional material that can be recorded here consists of 39
from Bedretto (Tessin), vii-viii, which were found in the collection of the
Naturhistorisches Museum at Basel among specimens of L. tenebricola (Wider) and
published under that name by Schenkel (1929: 10, p.p.).

The available material is listed here for completeness sake (all preserved in
NMB).

Wallis: 14 , Bez. Goms, Weg vom Hotel Jungfrau zum Märjelensee, 2190-2390
m, 21.vii.1923, E. Schenkel (lectotype); 19, Weg vom Hotel Jungfrau zum
Eggishorn, oberer Teil, 2400-2934 m, 7. vii.1924, E. Schenkel (paralectotype). 19
lg, Umgebung Saas Fee, vii-viii; 29 13, linker und rechter Talhang bei [Saas]
Almagel, vii., figured.

Tessin: 14 , Bedretto, rechter Talhang, vii; 39, Bedretto, vii-vili (L. tenebricola,
Schenkel, 1929, p.p.).

Lepthyphantes jacksonoides Van Helsdingen, spec. nov.
(Fig. 43-46, 66)

Lepthyphantes nigriventris; Schmölzer, 1962: 269 (Austria, Nordtirol).
Lepthyphantes spec. nahe jacksoni; Palmgren, 1973: 31, Fig. 2 (Austria: Nordtirol, Kärnten, Salzburg).

This new species closely resembles L. jacksoni Schenkel, from which it can be
distinguished by the shape of the lamella in the male palp, the stridulating files,
and (tentative conclusion) by its distribution.

Types. — ¢ holotype from Austria, Osttirol, Umbaltal W. of Prägraten, 1300-
1550 m, 4.viii.1973, P. J. van Helsdingen, from herbage along river; 29 23,
paratypes, collected with the holotype (whole series in ML).

Description. — Measurements (in mm). Total length, Q 2.15-3.2, 3 2.2-2.9;
length cephalothorax, 9 1.0-1.22, 3 1.0-1.25. Legs: Fe I 1.1-1.25 times length
cephalothorax in females, 1.1-1.3 times in males.

Coloration. Cephalothorax grey-brown, blackish-grey along margins. Legs light
brown. Abdomen with light dorsal and lateral surfaces, not unlike tenuis, at first

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes

Fig. 43—46. Lepthyphantes jacksonoides Van Helsdingen, spec. nov. 43, radical section, ventral aspect;
44, paracymbium; 45, male palp, lateral aspect; 46, epigyne. 44—46, X 128; 43, X 265

sight; dark grey V-shaped markings narrow or interrupted in the middle and light
lateral stripe broadly connected with the light areas of the dorsal surface, lighter
areas strewn with many white blotches. In males pattern much reduced, but often
still visible.

Stridulating files (Fig. 66) rather fine, especially basad, ridges closer together
than in jacksoni.

Legs with the usual set of spines, as indicated for the genus. Position of
retrodorsal (basal) spine on tibia I 0.29-0.34, length of this spine 0.31-0.35 mm in
®,0.21-0.28 mm ing . Tm 10.20-0.26.

Male palp (Fig. 43-45). Resembling in general the palp of jacksoni but differing
in the following respect. Dorsal and ventral branches of the lamellar tip not
diverging (cf. Fig. 39), but dorsal branch slightly curved and roughly parallel to
ventral one. Length of cymbium 0.40-0.47 mm (jacksoni: 0.42-0.45).

36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

Epigyne (Fig. 46) hardly different from that of jacksoni, with the same kind of |

depression near the tip of the basal part of the scape. Width of basal part of scape
0.210-0.225, of inner scape 0.160-0.190, of posterior median plate 0.250-0.275 mm.

Distribution. — So far known to occur in Switzerland (Glarus, Graubünden)
and Austria (Vorarlberg, Nordtirol, Osttirol, Salzburg, Kärnten). A female
specimen from Bulgaria (in cave “Dupocheto” near Velingrad, distr. Pazerdjik) is
thought to belong to jacksonoides, indicating a much wider distribution, but
confirmation at the specific level has to wait until males become available from
that area. Collected above 1,300 m and up to 2,400 m. It appears that the webs are
(always ?) built in the vegetation at some distance from the soil (up to 50 cm),
which is quite unusual for species of the tenuis group. The webs are thus more
easily detected than in most other species of the group. Collecting dates range
from June to September.

Rich material was available from Austria, mainly Nordtirol (figured), but the
description of the species is largely based on the following specimens.

Switzerland: 19 23 , Glarus, Stausee Garichti near Schwanden, 1500-1600 m,
9.ix.1972, P. J. van Helsdingen, in herbage (ML). 29 13 (palp only), Grisons (=
Graubünden), Dr. Carl (MHNG) (not included in the type-series).

Austria: 14 , Nordtirol, Zillertaler Alpen, Höllensteinloch, 1949, H. Janetschek
(NMB). 29 33 , Osttirol, Umbaltal W. of Prägraten, 1300-1550 m, 4.viii.1973, P. J.
van Helsdingen, from herbage along river (type-series; ML).

Lepthyphantes nigriventris (L. Koch)
(Fig. 47-49. 67)

Linyphia nigriventris L. Koch, 1879: 34, PI. 1 Fig. 22-22a (descr. 9, Siberia, Yenissey region).

Lepthyphantes nigriventris; Kulczynski, 1916: 20, Pl. 1 Fig. 29 (nigriventer; 2, Siberia). Ermolajev, 1934:
138 (West Siberia). Holm, 1945: 8 (Sweden), 54, Fig. 16 a-d (descr. 3 , genitalia depicted); 1951: 144
(Swedish Lapland); 1952: 122 (Swedish Lapland); 1973: 71, 95, Fig. 73-74 (redescription of type-
material, lectotype selected). Brinck & Wingstrand, 1949: 27. Hackman, 1954: 5, 53 (Newfound-
land).

Types. — The original material of Koch consisted of 39 from the Yenissey
region, Siberia. A lectotype (from Krasnoyarsk) was selected by Holm (1973: 95).
All specimens are preserved in the Naturhistoriska Riksmuseet at Stockholm. We
have examined the lectotype and one paralectotype.

Schmolzer’s record (1962: 269) probably belongs to the closely related species
from the eastern Alps, L. jacksonoides spec. nov.

Diagnosis. — L. nigriventris belongs to the jacksoni subgroup, which is
characterized by the pit or depression in the scapus of the epigyne, and by the
laterally projecting lamellate structure of the terminal apophysis in the male palp.
In nigriventris this projection is relatively short, and the position of the tooth on the
paracymbium is also different from the other species of the subgroup.

Measurements (in mm). Total length, 9 2.9-3.4, 4 (one specimen only) 2.6;

|
|

|

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 37

length cephalothorax, 9 1.23-1.32, 4 1.22. Legs: Fe I 1.2-1.3 times length
cephalothorax in females, 1.33 times in male.

Stridulating files (Fig. 67) fine, ridges close together.

Male palp (Fig. 47, 48). Paracymbium with a single, large and slender tooth on

Fig. 47—49. Lepthyphantes nigriventris (L. Koch). 47, male palp, lateral aspect; 48, radical section,
ventral aspect; 49, epigyne. Fig. 50—51. L. camtschaticus Kulczynski. 50, epigyne (holotype); 51, epigyne
(specimen mentioned by Schenkel, 1930). X 128

38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

the proximal branch, close to the posterior margin, and not at the base of, and
hidden behind, the distal branch (cf. jacksoni and jacksonoides). Lamella with
diverging dorsal and ventral branches, both rather straight. Terminal apophysis
with a lateral blade-like projection which is rather short and rounded (cf. jacksoni
and jacksonoides). Embolus without denticles on the basal ridge (cf. jacksoni and
jacksonoides). Length of cymbium (of specimen from Torne Trask) 0.46 mm.

Epigyne (Fig. 49) with a distinctly pitted scape; basal part of scape gradually |
widening posterad; lateral lobes of inner scape rounded, inner scape about as wide |
as the basal part; posterior margin of basal part sclerotized, reminding of |
tenebricola. Posterior median plate distinct in the ventral aspect of the epigyne. |

Width of basal part of scape 0.180-0.200, of inner scape 0.165-0.200, of posterior
median plate 0.235-0.250 mm.

Distribution. — Bonnet summarizes the distribution of this species as Norway, |

Sweden, Siberia. It has been found on several occasions in Swedish Lapland

(Holm, 1945, 1951, 1952; Brinck & Wingstrand, 1949), while the original material

of L. Koch from Siberia (Yenissey valley) was redescribed by Holm (1973: 95). A
single female has been recorded from Newfoundland (Hackman, 1954). We have
re-examined the specimen (Zoological Museum, Helsinki) and agree with the
identification; the lighter areas of the abdomen are strewn with white blotches, a
feature we have not observed in other specimens. L. camtschaticus Kulczynski,
from Kamtchatka, might belong to this species, though the shape of the epigyne
(no males known) is slightly different from that of nigriventris (see also the general
discussion, p. 46).

According to Dr. T. Kronestedt of Stockholm (pers. comm.) the webs of this
species are, again, found in the vegetation at some distance above the ground level
(see also under jacksonoides), not close to the surface as in most other species of
the tenuis group.

Material examined. — We have seen the lectotype and one paralectotype from
the Yenissey region (Briochowskij Islands, Chantajskoj, Krasnoyarsk) (NRS).
From Swedish Lapland we received specimens from Mr. T. Kronestedt: Is 29,
Torne Lapmark, N. of Torne Trask Lake, south side of Lullehatjärro mountain,
20-27.viii.1969, T. Kronestedt (ML).

Canada: 19, Newfoundland, St. Barbe Isl., Docters Hill, 29.vi.1949, E. Palmen,
from litter (Zoological Museum, Helsinki).

The figures are based on the specimens from Swedish Lapland.

Lepthyphantes camtschaticus Kulczynski
(Fig. 50, 51)
Lepthyphantes camtschaticus Kulczynski, 1926: 33, 57, Pl. 2 Fig. 19 (descr. © ; Kamtchatka). Schenkel,
1930: 17, Fig. 6 (descr. 9; Kamtchatka). Holm, 1973: 95, Fig. 75 (compared with L. nigriventris).

Type. — The © holotype is preserved in the collection of the Zoological
Institute at Warszawa (examined); it originates from Klutschevskoje, Kamtchatka.

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 39

Diagnosis. — Lepthyphantes camtschaticus belongs to the jacksoni subgroup
according to the median pit in the scape of the epigyne (absent in the holotype!).
Only two specimens have ever been mentioned in the literature, both females.
From these specimens it appears that the shape of the scape is slightly more
prolonged than in L. nigriventris. It is difficult, at this moment, to assess the value
of this character, because eastern Siberia is so poorly collected and no male
specimens have become available as yet. It is not impossible that we are dealing
with one polytypic species, which occurs in northern Europe (nigriventris), Siberia
(nigriventris), Kamtchatka (camtschaticus), and the Nearctic region (Newfound-
land; nigriventris) (see also the general discussion, p. 46).

Measurements (in mm). Female. Total length 3.4, length cephalothorax 1.20-
1.37, Fe I 1.3 times as long as cephalothorax.

Stridulating files fine.

Epigyne (Fig. 50, 51). The distal end of the basal part of the scape is more
prolonged posterad, thus giving it a more slender appearance; however, the width
is the same as in nigriventris (0.20 mm). In the holotype, no pit is visible on the
basal part of the scape, but in the specimen from Achomtenbay (Schenkel, 1930) a
well-developed pit is present (Fig. 51).

Fig. 52—54. Lepthyphantes cristatus (Menge). 52, male palp, lateral aspect; 53, radical section, ventral
aspect; 54, epigyne. 52, 53, X 150; 54, X 220

40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

Distribution. — Kamtchatka. |
Material examined. — We have studied the © holotype from Kamtchatka, |
Klutschevskoje, 10.viii.1909 (Warszawa), and the single 9 specimen recorded by |
Schenkel (1930) from Achomtenbay, Kamtchatka (NRS). |

Lepthyphantes cristatus (Menge)
(Fig. 52-54, 68)

Bathyphantes cristatus Menge, 1866: 121, Pl. 22 Fig. 46 (descr. 9 3 ; Poland (Danzig)). |

Lepthyphantes cristatus; Miller & Kratochvil, 1948: 139, 140, Fig. 2 a-d (compared with L. pallidiventris). |
Locket & Millidge, 1953: 386, Fig. 230A, 232E (9 3 ). Wiehle, 1956: 192, Fig. 319-326 (9 4 ). Wan- |
less; 1971222726, PI 2 Pl 6 Fig (0): 1973-1298 Pl 1FE18233R1767E19 730)! |

Lepthyphantes cristatus pallidus Miller & Kratochvil, 1938: 238, 242, Fig. 3 (descr. © ; Czechoslovakia). |

Lepthyphantes pallidiventris Miller & Kratochvil, 1948: 139, 140, Fig. 2 e-h (nom. nov. pro L. pallidus |
Miller & Kratochvil, brought to species level; compared with L. cristatus; 9 and a ).

Remarks. — Miller & Kratochvil (1938) described a subspecies of cristatus from
Czechoslovakia, which they subsequently raised to species level and, obligatory,
renamed pallidiventris (1948). We have examined material kindly sent to us by Dr.
Miller, and we did not find any substantial differential characters, notwithstanding
the apparent differences which can be observed in the 1948 illustrations (Fig. 2,
a-d, cristatus, e-h, pallidiventris). We are, therefore, of the opinion that pallidiventris
is nothing but an extremely light-coloured form of cristatus. Specimens of this kind
are frequently found, together with normally coloured specimens of cristatus, in
Nordtirol, and a subspecific status is therefore out of the question. Consequently
L. pallidiventris is listed in this paper as a junior synonym of cristatus.

Diagnosis. — L. cristatus takes an isolated position in the tenuis group because
of the horned cymbium, the occurrence of several tibial spines in the male palp,
and the deviating chaetotaxy.

Measurements (in mm). Total length, 9 2.3-2.7, 4 2.2-2.4; length cephalotho-
rax, Q 0.95-1.05,4 1.02-1.25. Legs: Fe I 1.1-1.25 times length cephalothorax in ©,
1.2-1.3 times ing .

Stridulating files fine (Fig. 68). Posterior tibiae (III and IV) with a retrolateral
spine and thus differing from the other species of the tenuis group.

Male palp (Fig. 52, 53) at once distinguished from the other species of the group
by the presence of four spines on the tibia and small tubercles at the proximo-
lateral and proximo-mesal corners of the cymbium. Paracymbium with posterior
margin slightly convex posterad and bearing a narrow ridge there which ends in a
small sharp tooth at its dorsal limit; a serrate crest-like elevation at the base of the
distal branch. Radix as usual, with the exception of the Fickert’s gland, which
appears to be modified in this species: the gland is followed distally by a second
inflated, but less spherical, section of the spermduct. Distal section of lamella a
simple flat structure, which fans out from the narrow base. Basal section of
terminal apophysis strongly sclerotized and protruding. Basal section of embolus
with one or two slender, tooth-like protrusions. Length of cymbium 0.44-0.51 mm.

Epigyne (Fig. 54) with basal part of scape rather suddenly widening from narrow
“stem” to comparatively broad posterior section with rounded tips; lateral wings

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 41

of inner scape roundish and gradually excavated anterad. Posterior median plate
well visible but not much wider than basal part of scape. Width of basal part of
scape 0.175-0.200, of inner scape 0.135-0.150, of posterior median plate 0.200-0.215
mm.

Distribution. — Up to 1940, L. cristatus had been recorded from nearly all
European countries, with the exception of Greece, Portugal and Spain. European
U.S.S.R. formed the eastern boundary, the Balkans and northern Italy were the
southern limits. In the north-west there were records from Iceland and Greenland,
but Braendegärd (1958: 86) correctly pointed out that Jackson’s (1930: 654) record
of cristatus is to be found in an appendix on the Icelandic spider fauna at the end
of his paper on the spiders of Greenland. Greenland thus has to be removed from
the list of countries where the species has been found. Moreover, it appears now
that in the Icelandic list Jackson only cites the older record for Iceland by Simon
(1892: clxxvii) as do the other references to the occurrence on Iceland of cristatus.
In the Paris collection a single male specimen from Iceland could be located
which quite likely constitutes the material on which Simon’s record was based.
This specimen is not a cristatus but a zimmermanni and we therefore are inclined to
believe that L. cristatus has never been found in Iceland.

A survey of the literature after 1940 revealed records of many of the European
countries within the range indicated above, and also from northern Spain (Denis,
1962 b; Gerona).

The only cave records are by Drensky (1935: 98, 99, 102), from Yugoslavian
caves.

We have examined material from Holland (not common), France, Austria
(common), Czechoslovakia (common), and Yugoslavia. It is a species of the leaf
litter and herbaceous layers in deciduous forests; in the Alps up to 1700 m, but in
Scandinavia it is said to occur frequently in moors (Braun & Rabeler, 1969: 42;
Palmgren, 1975: 59).

The figures are based on specimens from France (Dép. de Eure).

SPECIES INQUIRENDAE

The following species are considered to belong to the tenuis group, though
their exact identity could not be established. It appeared either impossible to
locate the type-material, or the specimens were incomplete and, therefore,
unidentifiable.

Lepthyphantes foliatus Denis, 1945: 52, Fig. 1, Pl. 2 Fig. 25 (descr. 9, Algeria).

The short description of the single female specimen Denis had before him when
naming this species contains remarks on size and coloration. It is accompanied by
a figure of the abdomen and a very small one of the epigyne. The latter is strongly
suggestive of that of L. tenuis because of the anchor-shaped scape and the rounded
lateral lobes. The size of the specimen agrees with that of tenuis, and so does the
abdominal pattern, be it that in the latter the dark spots seem to be very
contrasting with the lighter surroundings. L. tenuis is known to occur in Algeria,

42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

and it is quite possible that Denis had this species in the small collection of spiders |
from that region he had received from the British Museum (Natural History), |
London. The only answer to this question would be a re-examination of the |
specimen, but neither Denis’ private collection, now at the Muséum National |
d'Histoire Naturelle at Paris, nor the collection at London seem to contain the 9 |

holotype of L. foliatus. Its loss during the war has been suggested.

Lepthyphantes suldalensis Strand, 1903: 10, 17, Fig. 2 a-b (descr. , Norway).

The illustrations and description of this species strongly suggest it to belong to |
the tenuis group, even though Strand himself compared it with Lepthyphantes
obscurus (Blackwall). The original specimens could be located in the Paris |
Museum. There are two specimens, a subadult ¢ and an adults , but in the latter |

most legs and both palps are lacking. Thus we are deprived of the most reliable set
of characters. The remnants of the adult specimen give us no decisive answer to
the question of the identity. It is not very likely that there occurs a separate
species in southern Norway and we tend to refer it to one of the other species of
the tenuis group. Judging by the size of the adult specimen (length cephalothorax
0.90 mm, length Fe I 1.12 mm) it could be mengei, flavipes, or tenuis (excluding
herbicola on zoogeographical grounds). However, the stridulating files and shape
of the chelicerae exclude flavipes (which has a latero-basal boss and the

stridulating files very coarse, if visible at all). Comparison with specimens of

mengei and tenuis suggest the former, mainly because of the fine stridulating files.
L. mengei is common in Scandinavia, while L. tenuis appears to be much rarer.

Lepthyphantes tenoides Kolosváry, 1938: 66, 74, Fig. b, c (descr. 9, Yugoslavia).

All that can be said at present is that the epigyne reminds of that of L. tenuis, the
very reason why it was called tenoides. It certainly belongs to the tenuis group, but
without the specimen it is impossible to place the species more precisely.

Nearctic representatives of the tenuis group

Beside L. tenuis (Alaska, British Columbia, Washington) and nigriventris (New-
foundland) there are a number of species which occur in the Nearctic Region and
probably belong to this species-group: cracens Zorsch, described from New York;
zebra Emerton, occurring in the eastern United States; zelatus Zorsch, closely
related to zebra and found in Washington; and zibus Zorsch, also from
Washington. The reader is referred to the treatise on the genus Lepthyphantes in
the United States by Zorsch (1937).

DISCUSSION

From the outset we intended to present a survey of the tenuis group of species:
to provide useful means for the recognition of the species, to summarize the
known distributions, and to erect a framework of reliable characters for
establishing infra-group relationships and for a correct placing of species to be
described in the future.

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 43

NGEI FLAVIPES
MEN

HERBICOLA

È TENEBRICOLOIDES DRENSKYI

vit

Fig. 55—60. Male chelicerae (but female of flavipes) of Lepthyphantes species, showing stridulating files.
SAP SLD, kody C0 6 NWSE Si 06 IBIS RSS 2.6 ers)

The approach had to be typological. The species treated here differ in a limited
number of usable, mainly genital characters, and the differences are often so
minute that variation is hard to trace. Apart from size and coloration, variation in
the used differential genital characters only became apparent in a few instances,
and then is discussed in the text. Most likely variability is as normal a phenomenon
here as in other taxa, but it has to be studied independently, using more advanced
methods such as scanning electron microscopy. In our present study it plays a
minor role.

44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

If one thing has become clear when studying this species-group, it is the poor
state of knowledge of the group as a whole. This certainly is the result of
inadequate literature, which, in the past, made it hard to identify a specimen at all. |
As pointed out in the introduction, we noticed a relatively high percentage of,
misidentifications in old collections. New species names appeared to have been
proposed for specimens which in fact belong to the commonest members of the |

|
|
|
|

ZIMMERMANNI SPINIGER FLORIANA

JACKSONOIDES

TENEBRICOLA JACKSONI

Fig. 61—66. Male chelicerae of Lepthyphantes species, showing stridulating files. X 128

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 45

tenuis group. All this certainly did not help to clarify the dark points in our
knowledge of the interspecific relationships, the diagnostic value of traditionally
used characters, and the distributional patterns. Consequently we have attempted
to improve upon the unsatisfactory situation. By doing so, we became much aware
of all the existing gaps, many of which we are unable to fill at present. That our
knowledge of the distribution of northern species such as L. nigriventris and
camtschaticus is fragmentary is anderstandable because of the inaccessibility of
these areas. But when we have to state that we really do not know anything certain
about the distribution of zimmermanni in the Balkan Peninsula, we have to admit
that this is the result of lack of interest rather than lack of opportunities. Even
though the Swiss and Austrian Alps are easily accessible and indeed have been the
field of activity of several arachnologists, we still lack insight in the distributions of
several species (again zimmermanni, among others), including their altitudinal
zonation. It was only recently that we discovered the existence of two alpine
species in that area, jacksoni in the west and jacksonoides in the east, without being
able to establish the boundary, or the zone of overlap, between the two as a result
of insufficient collecting of “common” Lepthyphantes species.

Characters used.

Hardly paying any attention to coloration and abdominal patters, which we
suspect to be variable, we have incorporated size limits, proportional leg lengths,
and chaetotaxy into the diagnosis. The chaetotaxy provide diagnostic characters in
two cases only, viz., in spiniger, where the metatarsi bear more than the usual,
single dorsal spine, and in cristatus, with its extra prolateral spines on tibiae III and
IV.

The chelicerae show constant dentition through the whole group. The
stridulating files, which are always present (with the flavipes < as the only
exception), have been used, more than before, to characterize species, though
only in a comparative, qualitative way. The importance of this character lies in the
assumption that it plays an active role in the maintenance of hybridization
barriers, effectively isolating the species from each other. The organ is used during
courtship, and we may assume that differently built stridulating files produce
different vibrations in the web, and that positive response of a female is only
evoked by a male of the same species, producing the right vibration. Comparative
use of the files in the present paper is made possible by presenting figures of the
male stridulating files of all species. A quantitative analysis has not been
undertaken because several technical problems are involved, such as the changing
interstices of the ridges of the files from base to apex. All epigynes are of
comparable structure, i.e. a long sigmoid scape with a stretcher on top, arising
from the anterior wall of an atrium, which at the posterior side is bordered by the
medially incised posterior median plate. There is no reason to assume basic
differences in the functional properties of the organ in the different species. Since
the inner scape, apart from the shape of the lateral wings, has not been included in
this study we cannot make use of these parts in differentiating the taxa. From
Wanless’ study (1973) of the epigynes of the British species of this genus it is clear
that not much can be gained from these hidden parts. Only cristatus is slightly

46 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

NIGRIVENTRIS CRISTATUS

= > ;
ee ohm

A

Fig. 67—69. Male chelicerae of Lepthyphantes species, showing stridulating files. 67, 68, X 128; 69,
X 135

different from the other species. We have observed the scape, therefore, only in

the ventral aspect, using the shapes of the basal part of the scape, the visible
lateral wings of the inner part, and, of course, the presence or absence of a pit or
depression on the basal part.

As to the latter, we may speculate about the functional properties of the pit and
depression. From the analysis of the functions of the different elements of the
male palp and the epigyne in L. leprosus (Ohlert) (Van Helsdingen, 1965), we know
that the male palp is firmly anchored on the epigyne by, among others, the male
paracymbium. We have already pointed out that in all species with pitted scapes
or scapes with depressions the paracymbium of the male palp bears a long tooth. It
is not impossible that the large tooth on the paracymbium is functionally
correlated with the pit or depression. We must keep in mind though, that the
larger tooth on the paracymbium is differently situated in the different species: at
the base, and rather at the inside of, the distal branch (tenebricola, jacksoni,
jacksonoides), or on the proximal branch (nigriventris).

In the male palp, the following characters have been used: dentition of
paracymbium, shape of lamella, dentition of basal section of embolus, and possible
conspicuous parts of the terminal apophysis. Of these the paracymbial characteris-
tics and the presence of a conspicuous protruding portion of the terminal
apophysis have been used for subgroup delimitation. The other characters could
not be used for this purpose, but only contributed in a purely typological way to
species differentiation.

Infra-group relationships.
With the aid of the characters mentioned, we can distinguish certain subgroups

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 47

or clusters of species within the tenuis group, though it is not possible to divide all
species among these clusters. Starting with the species with pit-bearing scapes, we
may bring together nigriventris (Fig. 49), camtschaticus (Fig. 51), and tenebricola
(Fig. 37).

The scape does not have a pit in the holotype of camtschaticus, but it is present
in a second specimen from the same region; camtschaticus also is suspected to be
very close to nigriventris, with which it might form a superspecies or polytypic
species. However that may be, both have the typical sclerotized posterior margin
of the basal part of the scape, a feature we know so well from tenebricola. This
latter species is now revealed as exceptionally possessing a pitted scape (Fig. 37).
The male palps in the two species mentioned here (we do not have a male of
camtschaticus) have one thing in common, viz., the relatively large, blade-like
projecting part of the terminal apophysis (Fig. 36, 47); in both the paracymbium
bears a strong tooth, though of different position and in the case of tenebricola
accompanied by a second tooth. In two other species, jacksoni and jacksonoides,
both male characters are equally present (Fig. 39, 40, 45, 46): the blade-like
projecting part of the terminal apophysis, which is even larger here and quite
conspicuous, while the paracymbium does have a large tooth, in position
comparable to the larger tooth of tenebricola. The females of these two species do
not have pitted scapes, but both have the rather conspicuous depression near the
tip of the exposed basal part of the scape (Fig. 42 and 46). The pit and the
depression found in this subgroup might be functionally analogous structures.

A second subgroup is constituted by L. zimmermanni and the closely resembling
spiniger and floriana. The epigynes of these three species are hardly different as to
the shape of their scapes; all three have proportionally broad posterior median
plates. The males have a number of features in common, but as a group they
cannot be clearly delimited from the other species. For instance, the basal section
of the embolus bears denticles in the three species mentioned, but so it does in
tenuis and drenskyi. Of the latter the female is still unknown, but the female of
tenuis differs from the zimmermanni type (compare Fig. 16 and 24, 30, and 34).
L. herbicola would, according to the epigyne (Fig. 15), fit in with the zimmermanni
type, but there are no denticles on the embolus, and the shape of the lamella
deviates considerably from what is found there (compare Fig. 13 and 27, 28, and
Bi).

The teeth on the paracymbium give no clue; they are quite useful for the
separation of the species, but relationships cannot be inferred from their numbers,
positions, — or absence. We have not succeeded, therefore, in delimiting the
zimmermanni subgroup more precisely.

We feel that cristatus stands quite isolated, forming a possible link with one of
the other (natural?) species-groups of the genus; the spinose palpal tibia, the
cymbium with its two tubercles, the spatulate lamella, and the strongly sclerotized
basal section of the embolus, are not found in any of the other species.

L. mengei and flavipes seem to be closely related, which might be a mere
suggestion because of their small size and the absence of denticles on the embolus,
which character also might be correlated with size. L. perseus from Iran, described
as new in this paper, most likely is related to these two species, even though the

48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

embolic dentition is present. L. tenebricoloides, endemic to Madeira and the
Canary Islands, rather strongly deviates from the general type; it is larger than
most other species, and lacks denticles on the embolus as well as on the
paracymbium. L. drenskyi and tenuis resemble each other very closely in the shape
of the lamella and the number of denticles on the basal section of the embolus.

Zoogeographical remarks and possible evolutionary lines.

If we correctly have brought together tenebricola, nigriventris, camtschaticus, —

jacksoni, and jacksonoides in one natural subgroup, we may try and find an |

explanation for the present distribution of the species. Of the species under |

consideration, tenebricola certainly has the widest and most central distribution. It

occurs in Europe and probably in Asia (Kamtchatka record), as far north as

Lapland, as far south as Spain and Italy. In the Alps it reaches an altitude of 2,000
m. In Swedish Lapland it occurs together with nigriventris, which may reach much
further north and is more adapted to open grassland habitats, while tenebricola can
be classified as ombrophilous. L. nigriventris may have evolved from northern
populations of tenebricola, which became isolated by the advancing ice-cover in
the colder pleistocene periods, and survived in the relatively mild and ice free
regions of southern Finland and Siberia (Yenissey region). A similar situation may
have occurred in the Alps, where jacksoni and jacksonoides have a subalpine to
alpine distribution (1,300 m and above) and are found in more open vegetation
outside the forest. L. tenebricola does not occur above 2,000 m and is more or less
restricted to the forested areas. Its distribution probably is limited by, or coincides
with, the timber-line. The northeast-southwest differentiation into two species
may result from the former existence of isolated refugia in the Alps. It is not likely
that they represent cases of Nunatak survival, because they are now not of
exclusive alpine distribution. We rather think of isolated regions, cut off from the
main population of tenebricola by chains of glaciated mountains. We know of many
of these areas that remained free of ice during the main glaciations (cf. Holdhaus,
1954, map 36).

As pointed out when summarizing the distribution of L. zimmermanni, this
species has a wide European distribution, which, however, leaves a remarkable
hiatus in the eastern Alps and the mountainous regions of the Balkans. All
specimens examined from Switzerland came from the western part, mainly the
northwestern plain. The species does not occur in the eastern part, nor in
Nordtirol, while in Czechoslovakia it is said to be very rare (Buchar, pers. comm).
From Hungary there is no record; from Rumania it was erroneously mentioned
and the specimens concerned are described as a new species (floriana) in this
paper. South of the Alps it is known to occur in Italy and northern Yugoslavia.
From Bulgaria there is an isolated cave record, but we must not forget, of course,
that the European population continues eastward in Poland and the European part
of the U.S.S.R. L. floriana in our opinion is closely related to zimmermanni; so far
it is only known from one locality near Bucarest and one in Bulgaria; it may have
evolved there from a marginal population of zimmermanni.

In the southwest we have a comparable case. Here we find L. spiniger, which, if
we are not mistaken, is restricted to the Pyrenees, where it inhabits the entrances

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 49

of caves and dolines, as well as the litter stratum of pine forest. There are many
records of zimmermanni from the Pyrenees (in Denis’ papers on the spider fauna of
that region) and at present the two species are not clearly isolated geographically.
It is impossible to speculate here, with our restricted knowledge of the ecological
preferences of the two species and of the details of their distributions in the
Pyrenees, about the possible influence of the glaciations on the development of
these two closely related species.

Only two species have been reported with certainty from Madeira and the
Canary Islands, viz., tenuis and tenebricoloides. According to Schmidt (1975 a and
b) both species are subject to more than the usual variation in this archipelago. A
specimen of tenuis from La Palma was reported (Schmidt, 1975 b) to be much
darker than specimens of the mainland populations, a specimen from Gomera
appears to be much smaller (Schmidt, 1975 a). Specimens of tenebricoloides from
Gomera (Schmidt, 1975 a) show differences in the genitalia as well as in size, as
compared to the original series from Madeira. It is not surprising that Madeira and
the Canary Islands have their own endemic representative (tenebricoloides) of the
tenuis group. This is a well-known phenomenon in many groups of animals, and
there are many examples of endemic spiders from these islands. Future
investigations may bring to light to what extent the different islands of the
archipelago have their own characteristic populations of tenebricoloides or related
species. Several islands are not yet investigated. As to tenuis, we can hardly expect
to find much differentiation among the island populations. Again we do not know
exactly on which islands the species occurs (so far recorded from Madeira,
Gomera, La Palma, and also from the Acores). But tenuis apparently is easily
dispersed by man, as witness the occurrence in New Zealand, and it is not very
likely that different populations have evolved under these conditions. The region
certainly is worth special attention.

At the present stage nothing is to be gained from a speculation on the
relationships among the remaining species. Our insight in the meaning of the used
characters is too superficial. However, progress is helped by a number of factors,
which we hope to have made possible through this contribution. In the first place
many more records are needed for all the species of this group, with as many
particulars on habitats to be recorded as possible. For some species the interest
should be focussed on certain regions, viz., the Alps, the Carpathian Mountains,
the Balkan Mountains, and the Apennines. There is also no reason why the tenuis
group should not be represented in the Taunus or Caucasus. The occurrence of
tenuis in Afghanistan (Denis, 1958) and the recent discovery of perseus in Iran
clearly show that we may expect representatives of this species-group to occur
farther to the east and south. Our knowledge of the faunas of these regions is very
poor, certainly as regards the Linyphiidae. We kindly invite the attention of all
arachnologists to this interesting group of species.

ACKNOWLEDGEMENTS

For the loan of types and other material, for the gift of specimens, for general
information on species, and for other forms of cooperation, thanks are due to the
following persons and institutions: Dr. G. Alberti, Museo Civico di Storia

50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

Naturale, Trieste; Dr. J. Buchar, Karlovy University, Prague; Mrs. C. L.
Deeleman, Ossendrecht, Netherlands; Dr. C. D. Dondale, Entomology Research
Institute, Ottawa; Dr. E. Duffey, Monks Wood Experimental Station, Abbots
Ripton, England; Dr. R. R. Forster, Otago Museum, Dunedin, New Zealand; Dr.
M. W.R.de V. Graham, Hope Department of Zoology (Entomology), Oxford; Dr.
M. Grasshoff, Forschungsinstitut Senckenberg, Frankfurt (SMF); Dr. B. Hauser,
Museum d’Histoire Naturelle, Geneve (MNHG); Dr. M. Hubert, Muséum
National d’Histoire Naturelle, Paris (MP); Dr. T. Kronestedt, Naturhistoriska
Riksmuseet, Stockholm (NRS); Dr. C. Leonardi, Museo Civico di Storia Naturale,
Milano; Dr. A. Martelli, Museo Zoologico de “La Specola”, Firenze; Dr. F.
Miller, Brno; Mrs. F. Niculescu-Burlacu, Institutul de Biologie “Traian Savu-
lescu”, Bucarest; Dr. W. Starega, Instytut Zoologiczny P. A. N., Warsawa; Dr. E.
Sutter, Naturhistorisches Museum, Basel (NMB); Dr. F. R. Wanless, British
Museum (Natural History), London (BM). Material preserved at the Rijksmuseum
van Natuurljke Historie at Leiden is indicated by ML.

REFERENCES

Alberti, G., 1967. La collezione araneologica del Museo Civico di Storia Naturale di Trieste. Catalogo
dei tipi. — Atti Mus. Civ. Stor. Nat. Trieste 26: 25—28.

Berland, L., 1932. Voyage de MM. L. Chopard et A. Méquignon aux Acores (aoüt-septembre 1930). II.
Araignées. — Ann. Soc. Ent. France 101: 69—84.

Bertkau, P., 1890. Arachniden gesammelt vom 12. November 1888 bis zum 10. Mai 1889 in San Remo
von Prof. Dr. Oskar Schneider: 1—11.

Blackwall, J., 1852. A catalogue of British spiders, including remarks on their structure, functions, oe-
conomy and systematic arrangement. — Ann. Mag. Nat. Hist. (2) 9: 15—22.

———, 1854. Descriptions of some newly discovered species of Araneidea. — Ann. Mag. Nat. Hist. (2)
13: 173—180.

———, 1864. A history of the spiders of Great Britain and Ireland, Part 2: 175—384, PI. 13—29. — Ray
Society, London.

Bokor, E., 1922. [Arthropoden der ungarischen Grotten.] — Barlangkutatas 9: 1—22, 45-49,

Bonnet, P., 1957. Bibliographia Araneorum 2 (3): 1927—3026. — Toulouse.

Braendegard, J., 1932. 3. Araneae. In: Isländische Spinnentiere 1. Opiliones, Chernetes, Araneae. —
Goteborgs Kungl. Vetenskaps- Vitterhets-Samh. Handl., (B) 2 (7): 8—36, Fig. 1—10.

———, 1958. Araneida. In: The Zoology of Iceland 3 (54): 1—113, Fig. 1—88.

Bragg, P. D., & R. E. Leech, 1972. Additional records of spiders (Araneida) and harvestmen (Phalangi-
da) for British Columbia. — Journ. Ent. Soc. British Columbia 69: 67—71.

Braun, R., 1960. Neues zur Spinnenfauna des Rhein-Main-Gebietes und der Rheinpfalz. — Jahrb. Nas-
sauischen Ver. Naturk. 95: 27—89.

———, 1969. Zur Autökologie und Phänologie der Spinnen (Araneida) des Naturschutzgebietes
“Mainzer Sand’’. Gleichzeitig ein Beitrag zur Kenntnis der Thermophilie bei Spinnen. —
Mainzer Naturw. Arch. 8: 1—107.

Braun, R., & W. Rabeler, 1969. Zur Autökologie und Phänologie der Spinnenfauna des nordwestdeut-
schen Altmoränen-Gebiets. — Abh. Senckenb. Naturf. Ges. 522: 1—89.

Brignoli, P. M., 1971. Note su ragni cavernicoli Italiani (Araneae). — Fragm. Ent. 7: 121—229, Fig.
1—136.

Brinck, P., & K. G. Wingstrand, 1949. The mountain fauna of the Virihaure area in Swedish Lapland, 1.
— Kungl. Fysiogr. Sällsk. Handl., NF. 60 (2): 1—70.

———, 1951. Idem, 2. — Kungl. Fysiogr. Sällsk. Handl., N.F. 61: 1—173.

Caporiacco, L. di, 1936. Saggio sulla fauna aracnologica del Casentino, Val d’Arno Superiore e Alta Val
Tiberina. — Festschr. Strand 1: 326—369, Fig. 1—S.

———, 1949. Una piccola raccolta aracnologica dei monti di Calabria. — Atti Mus. Civ. Stor. Nat
Trieste 17: 132—136, one Fig.

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 51

———, 1950. Aracnidi cavernicoli Liguri. — Ann. Mus. Civ. Stor. Nat. Genova 64: 101—110, Fig.
(SSD

Chyzer, C., & V. Kulczynski, 1894. Araneae Hungariae 2 (1), Theridioidae: 1—151, PI. 1—5.

Cloudsley-Thompson, J. L., 1948. Note on Arachnida 6. Spider records from Britain and Iceland. —
Ent. Month. Mag. 84: 240.

Dalmas, R. de, 1917. Araignées de Nouvelle Zélande. — Ann. Soc. Ent. France 86: 317—-430, Fig.
1—76.

———, 1922. Catalogue des araignées récoltées par le Marquis G. Doria dans l’ile Giglio (Archipel
Toscan). — Ann. Mus. Civ. Stor. Nat. Genova 50: 79—96.

Deltshev, C., 1972 a. A contribution to the study of spiders (Araneae) from the caves in Bulgaria II.
Genus Lepthyphantes in Bulgarian caves. — Bull. Inst. Zool. Mus. 36: 137—147, Fig. 1—10.

———, 1972 b. A review of spiders (Araneae) from Bulgarian caves. — Proc. 5th Int. Congr. Arachn.,
Brno 1971: 99—104.

———, 1973. A contribution to the study (Araneae) of spiders from the caves of Bulgaria. III. Ecologi-
cal notes on spiders (Araneae) from the entrance parts of the caves. — Bull. Inst. Zool. Mus.
38: 3947, Fig. 1, Tab. 1—2.

Denis, J., 1933 a. Chasses arachnologiques dans les Pyrénées-Orientales (Region de Banyuls-sur-Mer et
Vallespir). — Bull. Soc. Hist. Nat. Toulouse 65: 529—591, Fig. 1—4.

———, 1933 b. Liste d’araignees recueillies dans le département du Var. — Ann. Soc. Hist. Nat. Tou-
lon (1932-1933) 17: 85— 102.

———, 1934. Elements d’une faune arachnologique de l’ile de Port-Cros (Var). — Ann. Soc. Hist. Nat.
Toulon 18: 136—158.

———, 1935. Liste des araignées recueillies dans le département du Var (2e note). — Ann. Soc. Hist.
Nat. Toulon 19: 104—113.

———, 1938. A contribution to the knowledge of the spiderfauna of the Andorra valleys. — Proc. Zool.
Soc. London 107B: 565—595, PI. 1, Fig. 1.

———, 1941. Les araignées des Iles Canaries. — Ann. Soc. Ent. France 110: 105—130, Fig. 1—16.

—— —, 1945. Descriptions d’araignées Nord-Africaines. — Bull. Soc. Hist. Nat. Toulouse 79: 41—57,
Fig. 1—2, Pl. 1—2.

———, 1949. Sur quelques araignées de Provence. — Bull. Soc. Zool. France 74: 16—18, Fig. 1—4.

———, 1952. Etudes Biospéologiques 35 (1). Araignées récoltées en Roumanie par Robert Leruth,
avec un appendice sur quelques araignees cavernicoles de Belgique. — Bull. Inst. Roy. Sci.
Nat. Belgique 28 (12): 1—50, Fig. 1—47.

———, 1957. Araignées du massif calcaire des Eaux-Bonnes (Basses-Pyrénées). — Bull. Soc. Hist. Nat.
Toulouse 92: 245—258, Fig. 1—8.

———, 1958. Araignées (Araneidea) de l’Afghanistan. I. The 3rd Danish Expedition to Central Asia.
Zoological results 22. — Vidensk. Medd. Dansk Naturh. Foren. 120: 81—120, Fig. 1—54.

———, 1959. Quelques araignées cavernicoles des Pyrénées. — Ann. Spéléol. 14: 219—231, Fig. 1—5.

———, 1962 a. Les araignées de l’archipel de Madère (Mission du Professeur Vandel). — Anais Fac.
Cienc. Porto 44: 1—119, Pl. 1—12.

———, 1962 b. Quelques araignées d’Espagne centrale et septentrionale et remarques synonymiques.
— Bul. Soc. Hist. Nat. Toulouse 97: 276—292, Fig. 1—15.

———, 1963. Spiders from the Madeira and Salvage Islands. — Bol. Mus. Mun. Funchal 17: 29—48,

Fig. 1—12.

———, 1964. Spiders from the Azores and Madeira. — Bol. Mus. Mun. Funchal 18: 68—102, Fig.
1—23.

———, 1967. Quelques captures d’araignees Pyrénéennes (III). — Bull. Soc. Hist. Nat. Toulouse 103:
32—50.

Drensky, P., 1931. Paiatzi ot pescherite na Beulgaria. Höhlen-Spinnen aus Bulgarien. — Spis. beulg.
Akad. Naouk 44: 1—50.

———, 1935. Uber die von Dr. Stanko Karaman in Jugoslavien und besonders in Mazedonien gesam-
melten Spinnen (Araneae). — Izw. tzar. prirod. Inst. Sofia 8: 97—110, 7 Fig.

Dresco, E., 1949. Recherches souterraines dans les Pyrénées Centrales (Notes biospéologiques; année
1947). — Bull. Soc. Hist. Nat. Toulouse 84: 182—196.

———, 1960. Catalogue raisonné des araignées et des opilions des grottes du canton du Tessin (Suisse).

52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977 |

— Ann. Spéiéol. 14: 359-390, Fig. 1—26.

Dresco, E., & J. F. Jézéquel, 1961. Attribution d’un rang spécifique à Lepthyphantes zimmermanni Bert- |
kau 1890 s. sp. spiniger E. Simon 1929 (Araneae, Linyphiidae). — Bull. Mus. Nat. Hist. Nat. (2) |
33: 105— 108, Fig. 1—6.

Ermolajev, W., 1934. Materialien zur Spumeniaunea West-siberiens. — Fol. Zool. Hydrobiol. 7
130—148.

Fage, L., 1931. Araneae, Se serie, précédée d’un essai sur l’évolution souterraine et son déterminisme. |
Biospeologica LV’. — Arch. zool. Exper. 71: 91—291. |

Forcart, L., 1961. Katalog der Typusexemplare in der Arachnida-Sammlung des Naturhistorischen |
Museums zu Basel: Scorpionidea, Pseudoscorpionidea, Solifuga, Opilionidea und Araneida, |
— Verhandl. Naturf. Ges. Basel 72: 47—87. |

Galiano, E. F., 1910. Datos para el conocimiento de la distribucion geografica de los Aracnidos en Es- |
paña. — Mem. Soc. Esp. Hist. Nat. 6: 343—424. |

Hackman, W., 1951. Bidrag till kännedomen om spindelfauna i provinsen Tavastia borealis. — Mem.
Soc. Fauna Flora Fennica 26: 82—85.

———, 1954. The spiders of Newfoundland. — Acta Zool. Fennica 79: 1—99, Fig. 1—121, Map 1—S.

Hasselt, A. W. M. van, 1885. Catalogus Aranearum hucusque in Hollandia inventarum. — Tijdschr. —
Ent. 28: 113—188.

Helsdingen, P. J. van, 1963. The Micryphantidae and Linyphiidae (Araneida) of the Netherlands, with
some notes on the genus Lepthyphantes Menge, 1866. — Zool. Verh. 62: 1—38, Fig. 1—6.

———, 1965. Sexual behaviour of Lepthyphantes leprosus (Ohlert) (Araneida, Linyphiidae), with notes
on the function of the genital organs. — Zool. Med. 41: 15—42, Fig. 1—19, PI. 1—2.

Holdhaus, K., 1954. Die Spuren der Eiszeit in der Tierwelt Europas. — Abh. Zool.-Bot. Ges. Wien 18:
1—493, Pl. 1—52, one map.

Holm, A., 1945. Zur Kenntnis der Spinnenfauna des Torneträskgebietes. — Ark. Zool. 36A (15): 1—80,
Fig. 1—26.

———, 1951. Araneae. In: P. Brinck & K. G. Wingstrand, The mountain fauna of the Virihaure area in
Swedish Lapland. II. Special account. — Kungl. Fysiogr. Sällsk. Handl., NF. 61: 138—149,
Fig. 7—8.

———, 1952. Studien über die Spinnenfauna des Torneträskgebietes. — Zool. Bidr. Uppsala 29:
103—213, Fig. 1—26, Tab. 1—5.

———, 1959. Spindlar fran Harjedalens fjälltrakter. — Ent. Tidskr. 79: 216—230.

———, 1970. Notes on spiders collected by the “Vega” Expedition 1878—1880. — Ent. Scand. |
188—208, Fig. 1—42.

———, 1973. On the spiders collected during the Swedish expedition to Novaya Zemlya and Yenisey in
1875 and 1876. — Zool. Scripta 2: 71—110, Fig. 1—117.

Huhta, V., 1965. Ecology of spiders in the soil and litter of Finnish forests. — Ann. Zool. Fennici 2:
260—308, Fig. 1—14.

———, 1971. Succession in the spider communities of the forest floor after clear-cutting and prescri-
bed burning. — Ann. Zool. Fennici 8: 483542, Fig. 1—37, Tab. 1—16.

Hull, J. E., 1933. Nomenclature of British Linyphiid spiders: A brief examination of Simon’s French ca-
talogue. — Trans. North. Nat. Un. 1: 104—110.

Hull, J. E., & J. W. H. Harrison, 1937. The natural history of the Island of Raasay and of the adjacent
isles of South Rona, Scalpay, Longay, and Fladday. — Scott. Natural. 226: 107—113.

Jackson, A. R., 1930. Results of the Oxford University Expedition to Greenland, 1928. Araneae and
Opiliones collected by Major R. W. G. Hingston; with some notes on Icelandic spiders. —
Ann. Mag. Nat. Hist. (10) 6: 639—656, PI. 17.

Keyserling, E., 1886. Die Spinnen Amerikas. 2. Theridiidae, 2e Hälfte: 1—295, Pl. 11—21. — Nürnberg.

Koch, L., 1879. Arachniden aus Siberien und Novaja Semlja, eingesammelt von der schwedischen
Expedition im Jahre 1875. — Kungl. Svenska Vet.-Akad. Handl. 16 (5): 1-136, Pl. 1—7.

Kolosväry, G., 1928. Die Spinnenfauna der ungarischen Höhlen. — Mitt. Höhl. Karstforsch. 1928:

109—113.
———, 1938. Sulla fauna aracnologica della Jugoslavia. — Rasseg. Faun. 5: 61—81.
Kulezynski, V., 1887. [Symbola ad faunam Arachnoidarum Tirolensem.] — Rozpr. spraw. wydz. mat.-

przyrod. Akad. umiej. 16: 245—356, PI. 5—8.

VAN HELSDINGEN, THALER & DELTSHEV: Lepthyphantes 53

| ———, 1916. Araneae Sibiriae Occidentalis Arcticae. — Mem. Acad. Imp. Sci. (Petrograd) (8) 28 (11):
1—2 + 1—44, Pl. 14-15.
———, 1926. Arachnoidea camtschadalica. — Ann. Mus. Zool. 27: 29—72, PI. 2—3.
| Lehtinen, P. T., & A. Kleemola, 1962. Studies on the spider fauna of the southwestern archipelago of
| Finland. I. — Arch. Soc. Zool. Bot. Fennicae “Vanamo” 16: 97—114, Fig. 1—17.
| Leruth, R., 1935. Exploration biologique des cavernes de la Belgique et du Limbourg hollandais, xxviie
| contribution: Arachnida. — Bull. Mus. Roy. Hist. Nat. Belgique 11 (39): 1—34.
| ———, 1939. La biologie du domaine souterrain et la faune cavernicole de la Belgique. — Mém. Mus.
Hist. Nat. Belgique 87: 1—506, Fig. 1—61.
Lindroth, C. H., 1957. The faunal connections between Europe and North America: 1—344, Fig. 1—61,
| Tab. 1—9, Diagr. 1—11. — Stockholm.
| Locket, G. H., & A. F. Millidge, 1953. British Spiders 2: 1—449, Fig. 1—254. — Ray Society, London.
| Locket, G. H., A. F. Millidge & P. J. van Helsdingen, 1970. An observation on the type material of
Lepthyphantes tenebricola (Wider) (Linyphia tenebricola Wider 1834). — Bull. Brit. Arachn. Soc.
| 1: 90—91, Fig. 1-8.
| Menge, A., 1866. Preussische Spinnen: 1—560 + 1—11, PI. 1—91. — Danzig.
} Miller, F., 1947. Pavouci zvifena hadcovych stepi u Mohelna. In: Mohelno, 7: 1— 107, Pl. 1—16.
| Miller, F., & J. Kratochvil, 1938. [Quelques nouvelles araignées pour la faune tchécoslovaque.] — Cas.
. Narodn. Mus. Praze 1934: 234—244, Fig. 1—4.
| ———, 1948. [Notes sur trois èspèces du genre Lepthyphantes Menge.] — Ent. Listy 11: 137—140, Fig.
| 1—2.
| Moritz, M., 1968. Lepthyphantes zebrinus (Menge), ein Wiederfund für die deutsche Spinnenfauna (Ara-
N neae, Linyphiidae). — Deutsch. Ent. Zeitschr., N.F. 15: 399—407, Fig. 1—12.
Niculescu-Burlacu, F., 1968. Contributii la studiul faunei de Aranee din Pädurea Bränesti. — Stud.
Cerc. Biol., Zool. 20: 89—94, Fig. 1—7.
| Palmgren, P., 1972. Studies on the spider populations of the surroundings of the Tvarminne Zoological
| Station, Finland. — Comment. Biol. 52: 1—133, Fig. 1—16, Pl. Fig. 1—23.
| ———, 1973. Beiträge zur Kenntnis der Spinnenfauna der Ostalpen. — Comment. Biol. 71: 1—52, Fig.
1—3.
EE 1975. Die Spinnenfauna Finlands und Ostfennoskandiens VI. Linyphiidae 1. — Fauna Fennica
| 28: 1—102, Fig. 1—23, Map 1—21.
| Polenec, A., 1958. Ekolska raziskovanja arahnidske favne iz nekaterih gozdnih tipov v okolici Kranja.
— Biol. Vestnik 6: 92—117.
| Roewer, C. F., 1942. Katalog der Araneae von 1758 bis 1940. 1 Band: i-viii, I— 1040. — Bremen.
Schenkel, E., 1925. Beitrag zur Kenntnis der Schweizerischen Spinnenfauna. — Rev. Suisse Zool. 32:
| 253—318, Fig. 114.
| ———, 1927. Beitrag etc. III. Teil. Spinnen von Saas-Fee. — Rev. Suisse Zool. 34: 221—267, Fig.
| 1—21.
| ———, 1929. Beitrag etc. IV. Teil. Spinnen von Bedretto. — Rev. Suisse Zool. 36: 1—24, Fig. 1—6.
| ———, 1930. Die Araneiden der schwedischen Kamtchatka-Expedition 1920— 1922. — Ark. Zool. 21A
| (15): 1—33, Fig. 1—13.
| ———, 1933. Beitrag etc. V. Teil. Spinnen aus dem Saas-Tal (Wallis) und von der Gegend zwischen
| Trins und Flims (Graubünden). — Rev. Suisse Zool. 40: 11—29, Fig. 1—2.
| ———, 1938. Die Arthropodenfauna von Madeira nach den Ergebnissen der Reise von Prof. Dr. O.
Lundblad Juli-August 1935. IV. Araneae, Opiliones und Pseudoscorpiones. — Ark. Zool. 30 A
| (7): 1—42, Fig. 1—12, Pl. 1—4.
Schmidt, G. E. W., 1975a. Spinnen von Gomera. — Zool. Beitr., Berlin, N.F. 21: 219—231, Fig. 1—10.
— — —, 1975b. Spinnen von La Palma. — Zool. Beitr., Berlin, N.F. 21: 233—237, Fig. 1—2.
Schmölzer, K., 1962. Die Kleintierwelt der Nunatakker als Zeugen einer Eiszeitüberdauerung. — Mitt.
| Zool. Mus. Berlin 38: 171—400, Fig. 1—17.
Simon, E., 1884. Les Arachnides de France 5 (2): 181—420, Fig. 1—198, Pl. 26. — Paris.
———, 1892. [Liste d’Arachnides recueillis en Islande par M. C. Rabot.] — Ann. Soc. Ent. France 60,
| Bull.: clxxvi-clxxvii.
———, 1897. Arachnides recueillis à l’île Madère par M. A. Fauvel en 1896. — Bull. Soc. Ent. France
1897: 111—113.

54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 1, 1977

—_____, 1929. Les Arachnides de France 6 (3): 533-— 772, Fig. 813—1112. — Paris.

Six, G. A., 1858. Lijst van spinnen in de provincie Utrecht gevonden en gedetermineerd. In: J. A.
Herklots, Bouwstoffen voor eene fauna van Nederland 2: 292—302.

Strand, E., 1903. Theridiiden aus dem westlichen Norwegen. — Bergens Mus. Aarbog 6: 1—23, Fig.
1-3.

Thorell, T., 1856. Recensio critica Aranearum Suecicarum quas descripserunt Clerckius, Linnaeus, De
Geerus. — Nov. Acta Reg. Soc. Sci. Upsaliensis (3) 2 (1): 61—171.

Vilbaste, A., 1973. [On the structure and seasonal dynamics of the spider fauna of Estonian mesotro-
phic peat-land forests.] — Eesti NSV Teaduste Akadeemia Toimetised 22, Biol.: 210—225.

Wanless, F. R., 1971. The female genitalia of the spider genus Lepthyphantes (Linyphiidae). — Bull.
Brit. Arachn. Soc. 2: 20—28, PI. 1—7.

———, 1973. Idem, II. — Bull. Brit. Arachn. Soc. 2: 127—142, PI. 1—9, Fig. 24.

Wider, 1834. Arachniden. In: A. Reuss, Zoologische Miscellen. — Mus. Senckenb. 1: 195—276, PI.
1418.

Wiehle, H., 1956. Spinnentiere oder Arachnoidea (Araneae). 28. Familie Linyphiidae-Baldachin-
spinnen. — Tierw. Deutschl. 44: i-viii, 1—337, Fig. 1—551. — Jena.

Wolf, B., 1934— 1937. Animalium cavernarum catalogus 3: 1—918.

Zorsch, H. M., 1937. The spider genus Lepthyphantes in the United States. — Amer. Midland Natural.
18: 856— 898, Fig. 1—93.

INDEX

(synonyms in italics)

arctica 18 gallicus 8 taczanowskii 19
arcuatus 30 henricae 12 tenebricola 12,30, 32
aspromontis 17 herbicola 16 tenebricoloides 21
borealis 24 jacksoni 32, 34 tenoides 42
camtschaticus 38 jacksonoides 34 tenuis 17, 42
concinnella 8 mengei 8,12 tomskica 8
concinnus 8 nigriventris 34, 36, 42 trucidans 19
cracens 42 pallidiventris 40 zebra 42

cristatus 24, 40 pallidus 40 zebrinus 15
drenskyi 22 perseus 11 zelatus 42
‘falteronensis 17 pygmaea 15 zibus 42

flavipes 12 sanfilippoi 18 zimmermannii 12, 18, 23
floriana 28 spiniger 27

foliatus 41 suldalensis 42

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J. VAN DER VECHT. — Studies of Oriental Stenogastrinae (Hymenoptera Vespoi-
dea), p. 55—75, fig. 1—49, pl. 1—2.

| Tijdschrift voor Entomologie, deel 120, afl. 2 Gepubliceerd 15-II-1977 |

N
Pe

STUDIES OF ORIENTAL STENOGASTRINAE
(HYMENOPTERA VESPOIDEA)

by
J. VAN DER VECHT

Burgemeester Vermeerlaan 4, Putten, Netherlands

With 49 text-figures and two plates

This is the third of a series of studies on the Stenogastrinae, an isolated and in
many respects particularly interesting group of subsocial wasps. It is also the first
paper to deal with the species of the Oriental region, where the group is much
more numerous and diverse than in the Papuan subregion. Whereas only 17
species, belonging to two genera, are known from the latter area (Van der Vecht,
1972, 1975), the Oriental region harbours about 50 species belonging to five
genera, two of which are new and described in this paper.

It is my intention to discuss the phylogeny and zoogeography of these wasps
after the completion of the taxonomic revision. It appears desirable, however, to
present at this stage a few notes on the relationships and the status of this group as
a whole.

The first author to deal with this question was H. de Saussure (1853, vol. 2: 4),
who divided the “Vespides”, often called Diploptera!) or diplopterous wasps, into
““Masariens’’, “Euméniens” and “Vespiens’’. Concerning the genus “Ischnogaster”
(junior objective synonym of Stenogaster, the only valid genus-group name
available for these wasps before 1914) this author wrote: “Ce genre est, par tous
ses caracteres, entierement intermediaire entre les deux tribus [Eumeniens and
Vespiens, solitary and social wasps, resp.]. Ses moeurs seules obligent a le classer
dans les Guépes sociales.”

Since then there has been little agreement with regard to the question whether
the Diploptera should be treated as a single family, embracing three or more
subfamilies, or as a superfamily containing two (Borner, 1919) or three families,
which in the latter case agree with the groups distinguished by De Saussure. That
this question is of some importance in connection with the systematic position of
the Stenogastrinae, may be shown by some examples of classifications proposed or
used in the past.

1) The name “Diploptera”, originally proposed by Latreille, in Cuvier, 1817, Règne Anim. 3: 502, was
rejected by De Saussure, because the longitudinal folding of the fore wings when at rest does not occur
in certain ‘“Vespides”, whereas it is also seen in certain wasps not belonging to this group “(les
Leucospidiens, par exemple)” (De Saussure, 1853, vol. 1: XVI). Nevertheless I shall use this term here,
because it appears to be the only available unambiguous name for the ““Vespides”’ sensu De Saussure.

SI)

56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 2, 1977

Smith (1857) used the term Diploptera, gave the three divisions family rank, and
placed /schnogaster in the Vespidae. Dalla Torre (1894, 1904) followed the
classification as given by De Saussure. Bingham (1897) also treated /schnogaster as
a member of the family Vespidae s. str., but remarked that this genus seems to
form “a link between the Eumenidae and Vespidae both in structure and in
habits.” Ashmead (1902) placed the three families of Diploptera, together with 13
other families of Hymenoptera (including Trigonalidae, Bethylidae, Chrysididae,
Pompilidae, Mutillidae, etc.), in one superfamily Vespoidea; he created a new
subfamily Ischnogasterinae, containing the African genus /schnogasteroides Ma-
gretti!) in addition to the type-genus, and placed this in the Eumenidae.

Bequaert (1918) and Bradley (1922) provided a classification in which all the
diplopterous wasps are regarded as a single family (Vespidae), subdivided in 10 or
11 subfamilies, 2 or 3 of these corresponding with the former family Masaridae, 3
with the Eumenidae, and 4 with the Vespidae. By placing the Stenogastrinae in
between the two last mentioned groups, the problem of the relationships of this
subfamily remains completely open, since no choice between the viewpoints of De
Saussure and Ashmead is necessary.

Börner (1919) divided the Diploptera, which he called ““Vespina’’, into two
families, each with two subfamilies: Vespidae with Vespinae and Eumeninae, and
Masaridae with Masarinae and Celonitinae; he could not examine /schnogaster
and /schnogasteroides, which he regarded as social wasps, but suggested that these
might be closely related to the Eumeninae.

In more recent literature, the term Vespoidea is either used for a group of
several families, including the Diploptera (Imms, 1925 and later editions; Essig,
1942; Bradley, 1958; Brothers, 1975), or exclusively for the Diploptera. In the
latter case, the superfamily may either contain one family Vespidae with several
subfamilies (Brues & Melander, 1932; Muesebeck c.s., 1951) or the usual three
families (Richards, 1962). The latter classification is here of particular interest,
because it requires an answer to the question of the affinities of the Stenogastri-
nae. Richards decided to place them again in the family Vespidae s. str. (social
wasps), although it is evident from his enumeration of the characters of this group
(1. c.: 22), that they take a very isolated position there.

Since then, the further study of these wasps has brought to light that the
differences between the Stenogastrinae and the other social wasps (Polistinae and
Vespinae) are considerably greater than would appear from this enumeration.
Richards himself has already pointed out that the Stenogastrinae differ from all
other Vespoidea in having the small subcircular pronotal lobe at a point almost
halfway between the tegula and the fore coxa (Richards, 1972). This character,
together with some others (long pointed clypeus and long narrow mandibles lying
alongside it) led him to remark that the Stenogastrinae are very different from
other social wasps and might even have evolved independently from some
Eumenes-like, solitary ancestor (Richards, 1971).

Spradbery (1975: 317) compared the Stenogastrinae with the Eumenidae and

1) This is a small group of solitary wasps which are structurally very close to Eumenes; Ashmead’s error
was corrected by Meade-Waldo (1913: 54).

VAN DER VECHT: Oriental Stenogastrinae 57

other Vespidae for 17 “biological characteristics’’!) and concluded that “the
Stenogastrinae exhibit a number of unique features which suggest that they
originated from an early vespoid ancestor and diversified in the specialized
environment ofthe deeply shaded rain forest.”

In the course of my studies of the Stenogastrinae some other characters have
been noticed which lend support to these ideas.

(1) Glossa and paraglossa do not have sclerotized pads (acroglossal buttons) at
their tips?). The Stenogastrinae agree in this respect with certain primitive
Masaridae. The absence of such pads in the few Stenogastrinae known to him was
already noted by De Saussure (1852a: 22; 1853, vol. 2: 3, footnote 1).

(2) The occipital keel runs down ventrally to meet the hypostoma close to its
base (see Richards, 1962: 11, and this paper, Fig. 1 and 2). This condition is not
found in the Polistinae and Vespinae. But the keel forks in several Zethinae *) and
in the South American genus Plagiolabra, one branch going to the hypostomal keel
and one to the mandible base. In some American Zethus species (Z. cinerascens
Saussure, cristatus Fox, spinipes Say, and probably others) the mandibular branch
is almost or entirely absent, and the situation is then very similar to that found in
the Stenogastrinae.

(3) Mandibles projecting beyond tip of clypeus, at least in © of some genera
distinctly crossing when at rest (Van der Vecht, 1972, Fig. 1 and 14), tridentate in
Q, often more or less reduced ing (in Eumenidae and Vespidae the mandibles of
Q usually have 4 or 5 teeth).

(4) Clypeus of 9 pointed at apex (in 2 sometimes rounded), differing from
clypeus of many Polistinae (©) in absence of projecting lateral angles of anterior
margin.

(5) Antennal sockets far apart, close to centre of eye-emargination, separated
from upper marging of clypeus (often only weakly indicated) by long (Liosteno-
gaster) to very long (Stenogaster) supraclypeal area.

(6) Apart from some characters already mentioned previously (see Richards,
1962), the wings of the Stenogastrinae are distinguished by (a) the shape of the
marginal cell (marginal vein bluntly angled at end of first abscissa, where r meets
Rs, and then running in an almost straight line to wing margin) and (b) by the
presence of only two closed cells in the hind wing, the costa being more or less
distinctly reduced distally. The row of hamuli is often once or twice interrupted.

(7) The male genitalia, particularly aedeagus and volsella, are rather different
from those of other diplopterous wasps, but are perhaps most similar to those of
certain Zethinae.

(8) The gaster (metasoma) of the pupa is strongly bent at the articulation of

!) This author has overlooked (Table 2, line 2) that Raphiglossinae and Zethinae?) are known to use
plant materials for nest construction.

2) The statement about the occurrence of acroglossal buttons in the Vespidae (Richards, 1962: 22)
should be corrected by inserting ‘(except in the Stenogastrinae)” after the word “present”.

3) Richards (1962: 4) and Bohart & Stange (1965) have used the name Discoeliinae Thomson, 1874, for
this subfamily, but in accordance with article 11 (e) of the Code the name Zethinae Saussure, 1855,
originally proposed as ““Zethites” for a group containing the genera Calligaster, Zethus and Discoelius,
has priority.

58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 2, 1977

segments | (petiole) and 2. This phenomenon does not occur in the Polistinae with
petiolate abdomen (the body of the pupa of Belonogaster, for example, is perfectly
straight), but it is well known from several Eumenini (example: photograph of

pupae of Delta emarginatum (L.) in Bonelli, 1973). Less attention has beengivento
the pupae of the Zethinae, the other Eumenid group of cell-builders with petiolate |
gaster. Except for a figure of the pupa of Calligaster williamsi Bequaert (“Zethus …

cyanopterus’’) in Williams’ paper on the biology of Philippine wasps (1919: 162, Fig.
94), which clearly shows “its bent attitude”, I looked in vain for information on
this subject in some literature directly at hand. The only pupa figured in Bohart &

Stange’s (1965) revision of American Zethus seems to have the gaster very slightly —

curved, but the drawing is small and rather schematic.

Fortunately I could study the pupae from nests of some Zethus spp. collected in
Surinam by Mr. G. van Vreden (Z. prominens Fox, no. 133 Mus. Leiden, and Z.
miniatus Sauss., no. 123) and by myself (Z. binodis (F.), no. 226). The first-
mentioned species is an interesting exception to the rule that the members of the
subgenus Zethus nest in insect burrows in twigs or wood; the nest found by Van
Vreden at Afobaka in a hollow tree is an irregular cluster of more than 30 cells,
each measuring about 3 x 1.5 cm, built of vegetable material consisting mainly of
fragments of parts of inflorescences of an unidentified plant species, pasted
together with a resinous substance. A pupa from this nest has the gaster sharply
bent, the angle between the axis of the petiole and that of the remaining segments
being about 70°. In a pupa of Z. (Zethoides) miniatus this angle is about 50°, and in
one of Z. binodis, which belongs to the same subgenus but has a relatively longer
petiole, it is only 35°. The few available figures seem to suggest that the angle
becomes sharper with increasing length of the petiole, but much more information
is required for a reliable conclusion. It would be particularly interesting to have
data on the pupae of species which resemble Stenogastrine wasps more closely in
petiolar length, such as certain members of the groups of Z. fuscus (Perty), Z.
strigosus Saussure, and Z. montezuma Saussure.

However, it is already possible at this stage to establish that the pupae of certain
Zethinae are very similar to pupae of Stenogastrinae, as figured by Iwata (1967,
Parischnogaster spp., Figs. 6-9, angle of gaster 25-45°) and Spradbery (1975,
Stenogaster concinna, Fig. 9, angle about 50°).

Some provisional conclusions from the data now available are:

(a) The Stenogastrinae are so different from the Polistinae and the Vespinae,
both in morphological and in ethological characters, that a taxon consisting of
these three groups cannot be regarded as monophyletic.

(b) The Stenogastrinae are likely to have evolved’ from a solitary cell-building
ancestor with elongate gastral petiole.

(c) Several characters, including those mentioned above under 2, 3, 7 and 8,
suggest that this ancestor was more closely related to the Zethinae than to the
Eumenini.

(d) Data on geographic distribution are not in contradiction with this theory.
The present distribution and diversity of the Zethinae in the Old World indicate
that this subfamily has been represented here long enough to allow for the
radiation of the Stenogastrinae in the Indo-Australian area.

VAN DER VECHT: Oriental Stenogastrinae 59

The consequences of these conclusions for the classification of the Diploptera

should be further studied in the framework of a revision of this group by modern
methods, for which Brothers’ recent study of the Aculeate Hymenoptera (1975)
may serve as an example. Meanwhile, if one accepts the subdivision of the
Diploptera into three (or more) families, a choice must be made whether to treat
the Stenogastrinae as a separate family or as a subfamily ofthe Eumenidae. In my
opinion the latter solution would then be preferable. However, I agree with
Richards (in litt., July 1976) that the best course might perhaps be to treat the
diplopterous wasps again as one family with several subfamilies, approximately
those listed on p. 4 of his revision ofthe Masaridae.

Ww |

KEY TO THE GENERA OF THE STENOGASTRINAE

Propodeum posteriorly with a very short constricted neck; its extreme edge
slightly raised, seen in profile as forming a minute tooth above the gastral
muscle. Head (excluding clypeus and mandibles) subcircular, at most moder-
ately wider than high (13 : 10). Mandibles of 3 without teeth, rather abruptly
narrowed from the middle, the apex curved and acute. Clypeus of 3 flattened
to concave. Posterior margin of hind wing of 3 normal. — New Guinea and
somemeishbounnpaslandsse $6500 ncn 0 ade cao Stenogaster Guérin
Posterior edge of propodeum not raised. Head (excluding clypeus and
mandibles) as a rule much wider than high. Mandibles of 3 , if without teeth,
not abruptly narrowed. Clypeus of 2 more or less convex. .......... 2
All segments of maxillary palpi about equally long (if segment 2 is twice as long
as 3, gastral segment 2 is not distinctly petiolate and occipital carina ends
below at short distance from hypostomal carina). — Oriental Region. .. 3

— Segment 2 of maxillary palpi more than twice as long as 3. .......... 6

Mesoscutum shining, with sharply impressed prescutal sutures, running on
outer side of anterior yellow marks backwards to level of tegulae. Head
broadly oval, width about 1.4 to 1.5 times height of eye. Spiracles of gastral
segment 2 clearly visible in dorsal aspect (Fig. 30). — Malaya, Sumatra,
BornneonBalawan andllavase er me Metischnogaster gen. nov.
Mesoscutum without prescutal sutures, or with ill-defined and shallow sutures
on dull mesoscutum (in group of Parischnogaster). Spiracles of gastral segment
PnoGvsiblenntdorsalfaspeectug Are ik) WERE BISHER Be mer 4
Occipital carina joins hypostomal carina near base of proboscideal fossa (Fig.
2). Gastral petiole long, more than 1.5 times length of thorax; segment 2 as a
rule distinctly petiolate. Antennae rather strongly swollen beyond segment 6 or
7. Parameral spine of male genitalia dilated, forming a broad lamella with short
ADI CALS PIMs varese a Sera. MEE eID es ide ae 5
Occipital carina ending at short distance from hypostomal carina (Fig. 1).
Gastral petiole less than 1.5 times length of thorax; segment 2 gradually
narrowing towards base, not petiolate. Antennae only slightly clavate.
Parameral spine normal. (Mid tarsi of 3 not dilated, symmetrical.) — Burma
and Thailand to Celebes and Java. ........ Liostenogaster Van der Vecht

60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 2, 1977

5. Clypeus longer than shortest interocular distance (Fig. 38 and 43). Mouthparts —
long, protruding beyond tip of clypeus over a distance well exceeding length of.
clypeus. Mid tarsi of very slender, symmetrical. — Borneo and Sumatra. |

PATIL BO IE AS EN ET A AMR Holischnogaster gen . nov. |

— Clypeus rarely a little longer than shortest interocular distance. Mouthparts _
short, protruding less far beyond tip of clypeus. Mid tarsi of 3 partly dilated, at
least one segment asymmetrical. — India to Celebes. ................

NR ETNA NT NES Parischnogaster Schulthess

6. Scutellum strongly convex, with sharp median carina. Mandibles of g as a
rule without teeth on inner side or, rarely (in species from continental S. E.
Asia), with one very blunt and faintly indicated tooth. Proximal part of
hind wing of 4 with opaque greyish or brownish seam at posterior margin.
Length to apex of second gastral segment exceeding 16 mm. — Southern
India to CCI Des TN ae ee Eustenogaster Van der Vecht

— Scutellum more or less convex, but without median carina. Mandibles of à
with at least one distinct tooth on inner side. Hind wing of ¢ without dark
seam. Length to apex of second gastral segment at most 14 mm. — New
Guinea and some neighbouring islands. .... Anischnogaster Van der Vecht

Metischnogaster gen. nov.
(Ei 9237 Pl?)

Head, including clypeus, wider than high (about 11:10 in 9, 12.3:10 ing),
and distinctly wider than thorax; width of head: height of eye = 14.5-15: 10 in 9
and 15.5-16: 10 in ). Clypeus moderately convex, shining and sparsely punctate
(Q) or practically impunctate (3 ); anterior portion bluntly keeled in the middle
and ending in a sharp point. Mandibles short, tridentate in 9, bidentate ing.
Segment 2 of maxillary palpi not lengthened; segment | of labial palpi long, but
shorter than 2-4 together. Antennae moderately clavate, segment 10 less than
twice as wide as segment 4; flagellum flattened and ventrally yellowish-white ing .
Vertex with impressed median line (sometimes partly indistinct) between anterior
ocellus and occipital carina.

Thoracic complex small, egg-shaped, and widest below tegulae; mesoscutum
with distinct prescutal sutures, running from anterior margin along outer side of
yellow marks to level of tegulae; parapsidal furrows starting a little in front of
anterior tangent of both tegulae, before reaching posterior margin of mesoscutum
a little dilated and bordered on outer side by fine carina. Scutellum moderately
convex, faintly carinate mid-anteriorly. Metanotum rather strongly convex in
profile. Propodeum shining, on posterior surface with oblique striation, transition
between posterodorsal and lateral areas rounded; median line impressed, most
distinctly at base and apex, and near the middle very slightly raised; valvulae
strongly reduced. — In the 4 the greater part of mesepisternum, metapleura and
sides of propodeum take part in the formation of a dull, densely pubescent, sunken
area. This area is bordered posteriorly by a sharp and more or less arcuate carina,
situated between posterior margin of metapleura, just below propodeal spiracle,
and apical margin of propodeum, close to posterior end of valvula.

VAN DER VECHT: Oriental Stenogastrinae 61

Fig. |—2. Rear view of head (mouthparts and clypeus omitted). 1, Liostenogaster nitidipennis (Saussure)

©, Borneo; 2, Metischnogaster drewseni (Saussure) 9, Java. Fig. 3. Wings of Metischnogaster drewseni

(Saussure) 2 , Java. Fig. 4—13. Submarginal cells 2 and 3 of right fore wing of Metischnogaster. 4—6, M.

cilipennis (Smith) 9, Malaya; 7—9, same, 2 , Malaya and Borneo (no. 9). 10, 11, M. drewseni (Saussure)

9, Java (in 11 transverse vein absent in both wings); 12, 13, same, , Java and Billiton. Scale lines re-
present 0.5 mm.

62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 2, 1977

a
1
1
1
\

ee

Fig. 14—20. Metischnogaster cilipennis (Smith) 9, Malaya. 14, head; 15, antenna: 16, mandible, lateral

and laterofrontal views; 17, maxillary palp; 18, labial palp: 19, gastral petiole, dorsal and lateral views:

20, gastral segments 2—6, lateral view. Fig. 21—27. Metischnogaster cilipennis (Smith) z , Malaya. 21,

head; 22, antenna; 23, left mandible, lateral view: 24, fore tibia and tarsus, and mid tibia; 25, paramere:
26, volsella; 27, aedeagus, lateral and dorsal views. Scale lines represent 0.5 mm.

VAN DER VECHT: Oriental Stenogastrinae 63

Legs slender, bearing numerous scattered long hairs, notably on coxae II and
III, on trochanters, at base of ventral side of femora and on outer side of tibiae and
tarsi; tibiae I of 9 on inner side with irregular row of short spines. Ing tibiae I
armed with a row of peculiar, slightly curved, spatulate setae (Fig. 24); tarsi II
symmetrical; segment 4 of tarsi I and II produced beneath into a long spine,
segment 5 with a shorter spine on ventral surface (Fig. 24 and 34).

Wings (Fig. 3-13) rather variable within the species as regards shape of second
and third submarginal cells and course of second recurrent vein, the latter
sometimes received in the third submarginal cell; sides of stigma almost parallel;
hind wing narrow, with posterior fringe of anal cell in 9 slightly lengthened, in z
very long.

Gastral petiole (Fig. 19 and 29) slender, greater part subcylindrical, somewhat
asymmetrically swollen at spiracles, apical two-fifths swollen into elongate bulbus
which is about 3.3 to 3.6 times as wide as the anterior part; tergite and sternite
almost completely fused, the suture separating them visible only on apical two-
thirds of ventral side of bulbus. Second segment (Fig. 20 and 30) at most shortly
petiolate; spiracles close to base on dorsal surface. Terminal segment conical (9)
or with flattened sternite (3 ). Male genitalia (Fig. 25-27, 35-37): parameral spine
long, with sharp tooth at base; volsellar digitus sickle-shaped; aedeagus com-
pressed, posterior part much higher than wide and near apex on each side with
small lamellar or spiniform process, ventral processes rudimentary.

Length to apex of second gastral segment: 9-11 mm.

Type-species: /schnogaster cilipennis Smith, 1857.

Distribution. — Malay Peninsula, Sumatra, Borneo, and Java.

Bionomics. — Our present knowledge of the life history of the two known
species (possibly the only ones existing) is entirely due to investigations made in
Malaya by H. T. Pagden (1958, 1962). The first paper deals with several
Stenogastrinae and contains, in addition to a description and some figures of the
peculiar characters of the 3 of “Stenogaster’’ cilipennis, a note on the habit of
hovering of these males and a drawing from a photograph of “the first nest ever
found” of this species. A closely related species, supposed to be undescribed, but
later identified as drewseni, was briefly mentioned. Further data are given in the
second paper, which is exclusively devoted to the habits of these two species.

The nest of M. cilipennis (Pl. 1) consists of an almost straight row of rarely more
than a dozen cylindrical cells, the first of which is usually attached to a hanging
thread of the fungus Marasmius (horse-hair blight) or to hanging rootlets; each
following cell is then built in a direct line with the preceding one, the openings of
the cells being directed sideward. The nests are protected by one or more “ant
guards”, constructed of the same material as the cells (probably made sticky); they
are difficult to find, being built in protected places, apparently often under
overhanging banks (above or close to water) and screened by vegetation. The nests
of the two species were found to be so similar, that no differences could be
detected. The construction of the nest takes a long time, one nest of M. drewseni
was observed to grow from 5 to 10 cells in about 4 months. Pagden noted that at
least two females and two males were present on this nest when the sixth cell was
about three-quarters complete. There is some evidence that the females of M.

64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 2, 1977

cilipennis collect food by hovering in front of spider’s webs from which they pick
small Diptera. The peculiar habit of hovering and patrolling in favoured shady
places was observed in the males of both species (PI. 2).

KEY TO SPECIES OF METISCHNOGASTER

Females

1. Interocular part of clypeus almost entirely yellow; yellow marks at inner orbits
coalescent with yellow clypeus or (in specimens from Borneo) separated.
Tibiae III yellow on outer side. Anterior (dark) part of dorsal surface of
propodeum indistinctly striate. Gastral petiole (Fig. 19) shorter in relation to
width of bulbus, with numerous long, outstanding, hairs on cylindrical part and
short hairs on bulbus. Gastral tergites 3 and 4 with basal yellow band (more or
less hidden under preceding tergites), which is interrupted medially and dilated
laterally sc ums een u ely a cilipennis (Smith)

— Interocular part of clypeus yellow with broad, dark brown, median stripe (Fig.
28); yellow marks at inner orbits separated from those on clypeus. Tibiae III
with short yellow line on outer side at base (absent in specimens from Java).
Dorsal surface of propodeum entirely obliquely striate. Gastral petiole (Fig.
29) longer and less hairy, the short hairs on bulbus much sparser. Gastral
tergites 3 and 4 each with two widely separated yellow spots. ...........

Pe RE ERR no vio drewseni (Saussure)

Males
1. Terminal three or four antennal segments mainly whitish dorsally (note that
the antennae may be more or less twisted). Markings of scutellum, metanotum
and propodeum ivory-white, the latter mark well separated from the carina
bordering the pubescent area. Metapleura without yellow spot. Gastral tergite
2 as arule without yellow spot beneath spiracle. Aedeagus on each side with
small subapical lamellar projection (Fig.27) ......... cilipennis (Smith)
— Terminal six or seven antennal segments (five ing from Palawan) mainly, or at
least partly, whitish dorsally. Markings of scutellum, metanotum and propo-
deum yellow (like the other thoracic markings), the last mentioned mark
abruptly dilated at apex, forming a band along posterior margin of shiny part of
propodeum. Metapleura with yellow spot beneath base of hind wing (not
always distinct). Gastral tergite 2 with elongate yellow spot beneath spiracle.
Aedeagus with small subapical tooth on each side (Fig.37) .............

Metischnogaster cilipennis (Smith) (Fig. 4-9, 14-27)

Ischnogaster cilipennis Smith, 1857, Cat. Hym. Brit. Mus. 5: 92,4 — “Malacca (Mount Ophir)”, leg. A.
Wallace (lectotype by present designation BM, no. 18.713); 1858, J. Proc. Linn. Soc. Zool. 2: 113
(Sarawak, Borneo); 1871, J. Proc. Linn. Soc. Zool. 11: 378 (cat.). — Dalla Torre, 1894, Cat. Hym. 9:

VAN DER VECHT: Oriental Stenogastrinae

Plate 1. Nest of Metischnogaster cilipennis (Smith), built on thread of horse-hair fungus (Marasmius), TA
cells below “ant guard”, with two female wasps (on cells 2 and 8) and one male (slightly reduced).
Malaya, Selangor, Ampang Reservoir, 9 May 1960.

Photograph by H. T. Pagden.

66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 2, 1977

Plate 2. Left: hovering males of Metischnogaster cilipennis (Smith) (A-E) and M. drewseni (Saussure) (F);
right: females of M. cilipennis searching at spider webs (slightly enlarged). Penang, Sungei Batu
Feringgi, 11 April 1960.

Photographs by H. T. Pagden.

VAN DER VECHT: Oriental Stenogastrinae 67

113 (cat.); 1904, Genera Insect. 19: 83 (cat.). — Schulthess, 1914, Zool. Jb. Syst. 37: 255 (in subgenus
Parischnogaster).

Parischnogaster cilipennis; Schulthess, 1927, Supplta ent. 16: 87 (cat.).

Stenogaster cilipennis; Dover, 1931, J. fed. Malay St. Mus. 16: 256 (Perak and Pahang, Malaya). — Pag-
den, 1958, Malay. Nat. J. 12: 141—145, Fig. 21, 22 (characters; bionomics in Malaya); 1962, Malay.
Nat. J. 16: 95-102, PI. 13-15 (bionomics in Penang). — Yoshikawa c. s., 1969, Nature & Life in S. E.
Asia 6: Pl. 34, 35 (nests, Malaya).

Note: for Ischnogaster cilipennis sensu Buysson, 1909, see under M. drewseni
(Saussure).

Material. — Malaya: Perak, 24 Jor Camp, 2000 ft, 22 Aug. 1922, E. Seimund
(BM); Pahang, 3 4 Sungei Tahang, 23 Nov. 1922, H. M. Pendlebury (BM);
Selangor, Ampang Reservoir, 5 9 (nrs. 981-983, 985, 987) and 1 3 (980) ex nest, 9
July 1947, 2 9 (1091, 1092) and 42 (1093-1096) ex nest, 5 Oct. 1947, H. T. Pagden
(BM, 3 9 22 ML); do. 1 9 29 Febr. 1948, ex nest, Comm. Inst. Ent. no. 10909,
with note “see also nest sent 28.4.48, G. N.” (BM); Selangor, 33 40 km N. of
Kuala Lumpur, 16 Sept. 1960, J. L. Gressitt (BISH, 13 ML), 13 Temple Park
near Kuala Lumpur, 14 June 1973, D. K. McAlpine (AMS); Malacca, 33 Mt.
Ophir (BM, lectotype and paralectotype; OUM, paralectotype).

Sumatra: N. E. Sumatra, 13 Serdang, Tandjong Morawa, Dr. B. Hagen (ML).

Borneo: Sarawak, | 9 “Sar.”, leg. A. Wallace (OUM), 13 Lundu, 21 Oct. 1909
(MCZ); East Borneo, 1 9 Tabang, Bengen River, 125 m, 5 Sept. 1956, 1 à
Balikpapan, Mentawir River, Oct. 1950, A. M. R. Wegner (ML).

Female

Head (Fig. 14), including clypeus, wider than long, preocular part of clypeus
about 1/5 of total length; interocular distance on vertex slightly longer than at
clypeus. Interocular part of clypeus in profile moderately convex, anterior part
almost straight; the latter with blunt median carina ending in a sharp point.
Mandibles: Fig. 16. Segment 2 of maxillary palpi slightly shorter than each of the
following segments, segments 4 and 5 produced beyond the implantation of the
next segment (Fig. 17). Segment 1 of labial palpi shorter than 2-4 together (4 : 5).
Antennal segment 3 slightly shorter than scape (7 : 8), segment 10 less than twice
as wide as segment 4. Ocelli large, transverse diameter of anterior one about 4/7 of
its distance from inner orbit, posterior ocelli smaller and almost circular, farther
from eyes than from each other (11 : 9), latter distance shorter than their diameter
BO).

Gastral petiole (Fig. 19) slightly more than 1.5 times as long as thorax, its length
about 6 times greatest width of bulbus.

Clypeus shining, slightly roughened medially, with scattered, superficial,
setigerous punctures. Front and supraclypeal area less shining, longitudinally
punctate-rugose, with short and low keel between and slightly in front of antennal
sockets. Vertex shining, with some shallow and ill-defined punctures. Thorax
generally smooth and shining, upper part of mesepisternum and lower part of
metapleura very finely and superficially striate or punctate-striate; mesoscutum
polished, with some scattered, setigerous, micropunctures. Dorsal surface of
propodeum finely obliquely striate, basally the striation partly obsolete, the striae

68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 2, 1977

meeting under a sharp angle medially. Gastral segments shining, almost smooth; a

fine microsculpture is most distinct on the third and following segments.

Short to moderately long hairs are present on most parts of the body, those on |
the face have in certain lights a golden shine; longer hairs are most conspicuous on |
mandibles, anterior part of clypeus, vertex, metanotum, propodeum, proximal part |

of legs, cylindrical part of gastral petiole and on hind margin of gastral sternites.

Ground colour of head and thorax dark brown to black, gaster and legs slightly |
paler; bright yellow markings: on face and mandibles as shown in Fig. 14, most of |
pronotal collar, interrupted band at hind margin of pronotum (bluntly angular on |
outer side at shoulders), two elongate spots at anterior margin of mesoscutum and |
two spots close to its posterior margin, two spots on scutellum, two on,
mesepisternum (ene close to base of fore wing, a lunate one below transverse |
suture), broad band on metanotum (incised posteriorly), large spot on upper part |
of metapleura, about posterior half of propodeum, coxae almost entirely, greater |
part of femora I, distal half or more of II and tip of III, tibiae I and II (except for |
brown stripe on inner side), dorsal stripe on tibiae III, basitarsus I above, short line |
on basitarsus II (sometimes lacking), curved stripe at lateral margin of tergal part |
of petiolar bulbus and some elongate spots on sternal part, more or less variable |
markings on postpetiolar part of gaster (Fig. 20). Wings subhyaline, strongly

iridescent, veins and stigma dark brown.
Length (h. + th. + t. 1 +2): 9.5-11 mm.

Male

and dilated (Fig. 24), outer (anterior) surface bare, flattened and rather dull

through fine sculpture, tarsal segments 1-3 of fore and mid legs with two rather |

long apical spines, 4 produced into a single long spine and 5 with shorter spine on

ventral side (Fig. 24); scattered hairs on tarsi much longer than in Q. Terminal

gastral sternite flattened, truncate at apex. Genitalia: Fig. 25-27.

Colour pattern conspicuously different from that of ©. Face dark brown (apex |
of clypeus paler) with broad, irregular, transverse yellow band (Fig. 21); ventral .

Head (Fig. 21) relatively wider than in the ©, anterior portion of clypeus |
shorter, mandibles weaker, bidentate, and with sharper teeth. Antennae flattened, |
apparently with rather soft underside (often shrivelled in dried specimens), scape |
relatively longer than in © (about 1.5 times as long as segment 3). Pubescent area |
on sides of thorax large, including entire mesepimeron. Femora I strongly curved |
in frontal view, tibiae I with row of spatulate setae (Fig. 24), tibiae II compressed

side of antennal scape and segment 2 polished and ivory-white, flagellum pale :

yellow beneath (on inner side when directed downwards), segments 11-13 mainly
yellowish white above (Fig. 22) (in from Sumatra segments 10-13). Thorax
without spots on lower part of mesepisternum and on metapleura; markings on

scutellum, metanotum and dorsal surface of propodeum together forming one |

broad, longitudinal, subrectangular, ivory-white stripe. Coxae II and III partly

brownish, femora brown with yellow mark at apex, tibiae I and II slightly darker |
than in ©, III brown with at most a short yellow line at base; tarsi brown, dorsal |
surface of basitarsus I yellowish. Gastral petiole dorsally with whitish stripe, |
dilated on bulbus (anterior part may be partly indistinct or absent); no lateral

VAN DER VECHT: Oriental Stenogastrinae 69

marks on bulbus and on tergite 2; 3 slightly paler anteriorly, 4 with basal yellow
band, dilated in the middle and at the sides, 5 with wider basal whitish band,
narrowed laterally, 6 mainly whitish, the dark posterior margin covering less than
one-third of the tergite, sternites 2-6 pale brown, each with two yellow spots;
terminal segment entirely whitish.

Length (h. + th. + t. 1 +2): 9-10 mm.

Metischnogaster drewseni (Saussure) (Fig. 2, 3, 10-13, 28-37)

Ischnogaster drewseni Saussure, 1857, Annls Soc. ent. France (3) 5: 315,4 — “Poulo-Pinang, Île de l’Ar-
chipel Indien”, leg. Drewsen (MHNG). — Smith, 1858, J. Proc. Linn. Soc. Zool. 2: 113 (Sarawak,
Borneo; Mt. Ophir, Malaya). — Dalla Torre, 1894, Cat. Hym. 9: 113 (cat.); 1904, Genera Insect. 19:
83 (cat.). — Buysson, 1909, Annali Mus. civ. Stor. nat. Genova 44: 312 (syn. of Ischnogaster cilipen-
nis). — Schulthess, 1927, Supplta ent. 16: 87 (cat., in synonymy of Parischnogaster cilipennis (Smith)).

Ischnogaster cilipennis; Buysson, 1909, Annali Mus. civ. Stor. nat. Genova 44: 312 (description of 9,
Lelemboli, Nias) [misidentification].

Stenogaster drewseni; Pagden, 1962, Malay. Nat. J. 16: 95-102, Pl. 13 (bionomics in Penang).

Material. — Malaya: Kedah, 2¢ near Jitra Catchment Area, 6 and 9 April 1928,
H. M. Pendlebury (BM, ML); Penang, 1 4 “Poulo-Pinang” (= Penang), leg.
Drewsen (type, MHNG), | 3 Batu Feringgi Catchment Area, 2 June 1960, H. T.
Pagden (BM), 1 9 14 Sungei Pinang, 1500 ft, 3 June 1962, ex nest of 15% cells on
Marasmius, H. T. Pagden (ML), 3g Batu Feringgi, 25 Febr. - 1 March 1963, M. A.
Lieftinck (ML); Selangor, | 4 Ampang Waterworks Reserve, 15 Aug. 1926, C.
Dover (BM).

Singapore: | 9 coll. Baker (USNM).

Sumatra: Nias Isl., 1 9 Lelemboli, Aug. 1886, leg. Modigliani (cilipennis det.
Buysson) (MCG); Palembang, 4 5 Pagar Alam, 750 m, 23 May 1935, Mrs. M. E.
Walsh (BM, 1g ML).

Belitung Isl: “West Billiton”, Tjerutjuk, sea level, 13 2 Aug. 1935, 1 © 24 14
Dec. 1936, F. J. Kuiper (ML).

Borneo: 13 “Borneo”, leg. F. Baczes (NMW); Sarawak, 1 3 “Sarawak”, 1897,
R. Shelford (BM); 8 @ Kuching, 1899-1900, Dayak collector, from R. Shelford
(OUM, 25 ML), 1 9 River Kapah, trib. of R. Tinjan (MCZ).

Palawan: 24 Brooke’s Pt., Tigoplan River, 500-700 ft, Philipp. Zool. Exp., F. G.
Werner leg. (CNHM, ML).

Java: West Java, Bogor (“Buitenzorg”), Djasinga, 150 m, at river Tjibarangbang,
23 April 1935 and Nov. 1936, 2 © Febr. and June 1937, M. A. Lieftinck (ML), 1
Q 24 July 1937, J. and E. van der Vecht (ML); 13 Wijnkoop Bay, Dec. 1936,
Mrs. M. E. Walsh (ML), 13 Djampang Wetan, Radjamandala, 1200 ft, Nov. 1937,
Mrs. M. E. Walsh (BM).

Female

Very similar to M. cilipennis, but easily distinguished by the characters
mentioned in the key on p. 64. Further differences are as follows.

Preocular part of clypeus slightly longer than one-fifth of length of head;
interocular part in profile slightly less convex than in M. cilipennis; in centre of
clypeus the interspaces between the punctures smooth and shining. Lateral areas

70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 2, 1977

of mesoscutum (on outer side of prescutal furrows) superficially rugose and |

slightly less shining than in M. cilipennis. |
Yellow markings rather constant, but the dark line on the clypeus frequently less |
narrowed at base than in Fig. 28.

Male

il
|
|
|

Apart from the colour characters mentioned in the key, distinguished by the mid |
tibia, which is less dilated (especially the basal half narrower), with outer side |

slightly convex and shiny and bearing a few outstanding hairs.

Yellow spot at apex of clypeus sometimes indistinct or absent; black supra-
antennal mark may be smaller than in Fig. 31; pale stripe on dorsal side of,
antennae (Fig. 33) usually present on the terminal six segments, rarely also on |
segment 7 (in one g from Java). Gastral tergite 1 with elongate dorsal spot on
bulbus, 2 with lateral spot below spiracle, 3 brown or with small lateral spot, 4 and |

5 with moderately wide yellow band in the middle of the tergite, separating an
anterior translucent area (whitish in living insect?) from the dark posterior part,

which is widened laterally, the yellow band often separated from a lateral spot of
the same colour, 6 whitish with rather narrow dark posterior margin; sternites 2-5 |
mainly as in ©, but some spots may be reduced or absent, 6 with irregular yellow |

band at base; terminal segment whitish, ventrally mainly yellow.

In the two males from Palawan only antennal segments 10-13 are mainly whitish |

dorsally, and the pale-coloured part of segment 9 covers less than its half. Since
the mid tibiae of these males are somewhat duller than in the other specimens,
examination of additional material from this island is desirable.

Holischnogaster gen. nov. (Fig. 38-49)

Head (Fig. 38 and 43), including clypeus, nearly as wide as high in 9, slightly |
wider than high in ¢ . Clypeus longer than wide, narrowed into a truncate point in |

Q, ing relatively shorter with nearly regularly triangular anterior portion.
Mandibles rather long, tridentate, crossing when at rest. Malar space short in 9,
slightly longer in ¢ . Mouthparts (Fig. 39 and 43) long, apical lobes of the narrow
glossa (in preserved specimens) not divergent; segment 2 of maxillary palpi longer
than other segments, but less than twice as long as any of these; segment | of labial
palpi about as long as 2-4 together. Antennae clavate, gradually swollen from
segment 6 to 10 (2) or 11 @ ), and here more than twice as thick as at segment 6.

Occipital carina meets hypostomal carina near base of proboscideal fossa. Vertex |

without impressed median line behind ocelli.

Thoracic complex (measured in lateral aspect) slightly longer than half the
length of gastral petiole (4 : 7), general shape as in Parischnogaster, but in 3 the
sides without any trace of specialized pubescent areas. Mesoscutum without
prescutal sutures.

Legs slender, not modified ing .

Wings mainly as in other Stenogastrinae, but the second submarginal cell only
moderately wide and narrowed towards the radial cell, the first cross-vein (Rs)
being rather strongly reclivous (front end farther from wing base than hind end).

VAN DER VECHT: Oriental Stenogastrinae TI

Nervulus (cu-a) usually placed before the fork of M and Cu; vein Cu,b strongly
reclivous. Hamuli (wing hooks) 6-8, as usual in this subfamily, widely and
somewhat irregularly spaced. Anal lobe of hind wing very small.

Gastral petiole (Fig. 40 and 41) slender, bulbus elongate, its width slightly more
than three times the smallest width of the cylindrical part. Spiracles of petiole and

Fig. 28-30. Metischnogaster drewseni (Saussure) 9, Penang. 28, head; 29, gastral petiole, dorsal and la-

teral views; 30, gastral segment 2, dorsal view. Fig. 31-37. Metischnogaster drewseni (Saussure) 3 „Java.

31, head: 32, mandible, lateral view; 33, antenna; 34, fore tarsus; 35, paramere; 36, volsella; 37, ae-
deagus, lateral and dorsal views. Scale lines represent 0.5 mm.

12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 2, 1977

Fig. 38—42. Holischnogaster gracilipes sp. n. 9, Borneo. 38, head; 39, mouthparts, ventral view; 40, gas-

tral segments | and 2, lateral view; 41, gaster, dorsal view, segments 3—6 extended to show yellow mar-

kings; 42, segments 2—6, ventral view. Fig. 43—49. Holischnogaster gracilipes sp. n. 2 , Borneo. 43, head

(length of glossa indicated by dotted lines); 44, gastral sternites 7—8, flattened; 45, paramere; 46, vol-

sella; 47—49, aedeagus, lateral, ventral and dorsal views, respectively. Scale lines represent 0.5 mm
(except Fig. 40 and 42).

VAN DER VECHT: Oriental Stenogastrinae 73

segment 2 visible in ventral aspect, the latter distinctly placed before midlength of
tergite. Segment 2 (Fig. 41) longer than wide at apex, slightly constricted at base.
Male genitalia mainly as in Parischnogaster mellyi group, but aedeagus more
strongly curved and higher.

Type species: Holischnogaster gracilipes spec. nov.

Distribution. — Borneo and Sumatra.

Note. — The type-species is closely related to Parischnogaster, as is evident from
the structure of the male genitalia. I have provisionally treated it as a separate
genus, because the lengthened mouthparts of both sexes and the non-dilated mid
tarsi of the male suggest that there may be important differences in behaviour with
regard to feeding and mating. Investigations on these points would be very
valuable. Also discovery of the hitherto unknown nest construction might give
important information on possible relationships.

Holischnogaster gracilipes spec. nov. (Fig. 38-49)

Material. — Borneo: Sabah, 3 9 53 Mt. Kinabalu, 5000 ft, 1-5 May 1973, K.
M. Guichard (holotype 9 and allotype g in coll. Giordani Soika, paratypes: coll.
Giordani Soika, 1 9 23 ML); 1& Penrissen, May 1899, Shelford leg. (paratype,
BM 1910-203); Sarawak, 1 © Mt. Dulit, primary forest, 26 Oct. 1932, Oxford Univ.
Exp., B. M. Hobby & A. W. Moore (BM 1933-254) (gaster lacking).

Sumatra: 13 Sungai Kumbang, Sept. 1915, Edw. Jacobson, no. 85-13 (antennae
and gaster lacking) (ML).

Female

Clypeus faintly shining, punctate on microsculptured, reticulate-coriaceous
ground; the punctures varying in size, small and close at the base, larger in centre
and anteriorly and here separated by one or more puncture widths; apex of
clypeus bluntly keeled. Supraclypeal area not distinctly separated from clypeus,
dull and densely punctate, at level of lower margin of antennal sockets with short
median keel (hardly more than elongate low tubercle). Frons and vertex dull, more
rugosely punctate.

Pronotum dull, transversely rugose-striate, the striae coarsest in anterior-lateral
angles. Mesoscutum and scutellum dull, densely rugose-punctate, the anteriorly
dilated space between median scutal lines superficially roughened and more
shining. Mesepisternum: upper part dull, irregularly obliquely striate, hypo-
epimeral area slightly more shining and superficially rugose, transverse suture with
row of punctures; ventral part swollen, rugosely punctate-striate, on each side with
trace of precoxal suture. Metanotum finely rugose-punctate, metapleura moder-
ately shining. Propodeum rather shining, finely obliquely striate, the striae sharply
defined on dorsal surface, somewhat irregular and running into puncturation at
lateral margins.

Gastral segments impunctate, with very fine and superficial microsculpture.

Face with short, appressed, golden pubescence and moderately long out-
standing hairs; longer hairs are present at apex of clypeus and on mandibles.
Pubescence of thorax and legs generally longer than in Parischnogaster mellyi
(Saussure), gastral petiole with rather long outstanding hairs.

74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 2, 1977

Ground colour dark brown to blackish; thorax, legs and abdomen partly pale
brown; the following parts yellow: mandibles (brownish at margins and on teeth), |
clypeus (except for narrow band at anterior margin and irregularly shaped mark in |
centre), underside of antennal scape, spot on genae at mandible base, pronotal |
collar, band at posterior margin of pronotum, narrowly interrupted medially and ‘|
strongly dilated at shoulders, two elongately triangular spots on anterior part of |
mesoscutum, broad band at base of scutellum and small spot on maxilla, large |
vertical spot on upper part of mesepisternum and an even larger one on lower |
part, bordering on the transverse suture, broad sublunate band on disk of |
metanotum and small triangular spot in anterior angle of lateral cavity, spot on |
metapleura beneath base of hind wing; propodeum except subquadrate area at |
base, produced on each side along anterior margin, and irregular band on lateral
surfaces; greater part of coxae, lines on femora and tibiae I, II, and distal half of
III, line on basitarsus of fore legs; markings on gaster as shown in Fig. 40-42. |

Length (h. + th. + t. 1 +2): 12-13 mm.

Male
Very similar to female, mainly differing in shape of head and clypeus (Fig. 43); |
yellow markings generally somewhat more extensive, but evidently rather |
variable; clypeus entirely yellow, supraclypeal area with yellow spot on each side |
(sometimes reduced, as in Fig. 43, or absent), yellow spot on genae larger than in |
Q, spots on mesoscutum longer, in some specimens produced backwards as an |
interrupted line which ends at level of posterior margin of tegulae; dark area at |
base of propodeum reduced, subtriangular; yellow line on tibiae III either entire
or narrowly interrupted; gastral tergite | with narrow lateral line on basal half of |
|

bulbus, 2 as in Q, 3 with broad yellow basal band (anteriorly with brownish area
narrowed towards the sides), the brown posterior part of the tergite covering less
than half the length in the middle and much less at the sides; 4-6 with broad
yellowish-white basal band, leaving only a narrow dark band at posterior margin, 7
yellowish-white on basal half; sternites mainly as in ©, terminal sternite with
yellow spot, its narrowed posterior part brownish. |

Length apparently not different from that of 9.

REFERENCES

Ashmead, W. H., 1902. Classification of the fossorial, predacious and parasitic wasps of the superfamily
Vespoidea, parts 6-8. — Canad. Ent. 34: 163—166, 203—210, 219—221.

Bequaert, J., 1918. A revision of the Vespidae of the Belgian Congo, etc. — Bull. Amer. Mus. nat. Hist.
39: 1—384, 6 pls.

Bingham, C. T., 1897. Fauna of Brit. India, incl. Ceylon and Burma. Hymenoptera I, Wasps and Bees,
579 pp., 4 pls. — London.

Borner, C., 1919. Stammesgeschichte der Hautflügler. — Biol. Zentralbl. 39: 145—186.

Bohart, R. M., & L. A. Stange, 1965. A revision of the genus Zethus Fabricius in the Western Hemi-
sphere. — Univ. Calif. Publs Ent. 40: 1—208, 354 figs.

Bonelli, B., 1973. Osservazioni eto-ecologiche sugli Imenotteri aculeati dell’ Etiopia, 5. De/ta emargina-
tum (L.). — Boll. Ist. Ent. Univ. Bologna 32: 27—46.

Bradley, J. C., 1922. The taxonomy of the Masarid wasps, including a monograph of the North Ameri-
can species. — Univ. Calif. Publ. Tech. Bull. 1: 369—464, 15 pls.

———, 1958. The phylogeny of the Hymenoptera. — Proc. X. Int. Congr. Ent. Montreal (1956) I:
265— 269.

VAN DER VECHT: Oriental Stenogastrinae 15

Brothers, D. J., 1975. Phylogeny and classification of the Aculeate Hymenoptera, with special referen-
ce to Mutillidae. — Kansas Univ. Sci. Bull. 50 (11): 483—648, 101 figs.

Brues, C. T., & A. T. Melander, 1932. Classification of Insects. — Bull. Mus. comp. Zool. Harv. 73:
1—672, 1121 figs.

Dalla Torre, K. W. von, 1894. Catalogus Hymenopterorum 9, Vespidae (Diploptera): 1—181. — Leip-
zig.

———, 1904. Hymenoptera, Fam. Vespidae. — In: P. Wytsman (ed.), Genera Insectorum 19: 1—108, 6
pls.

Essig, E. O., 1942. College Entomology, 900 pp., 308 figs. — New York.

Imms, A. D., 1946. A general textbook of entomology (6th ed.), 727 pp., 624 figs. — London.

Iwata, K., 1967. Report of the fundamental research on the biological control of insect pests in Thai-
land II. The report on the bionomics of Aculeate wasps — Bionomics of subsocial wasps of
Stenogastrinae (Hymenoptera, Vespidae). — Nature & Life in S. E. Asia, Tokyo 5: 259—293, 4
pls.

Jacobson, Edw., 1935. Aanteekeningen over Stenogastrinae. — Ent. Meded. Ned. Indie 1: 15—19, pl. 2,
fig. 2.

Meade-Waldo, G., 1913. New species of Diploptera in the collection of the British Museum. — Ann.
Mag. nat. Hist. (8) 11: 44—54.

Muesebeck, C. F. W., K. V. Krombein & H. K. Townes, 1951. Hymenoptera of America North of
Mexico, Synoptic Catalog. — Agric. Monogr. 2: 1—1420.

Pagden, H. T., 1958. Some Malayan social wasps. — Malay. Nat. J. 12: 131—148, 22 figs.

———, 1962. More about Stenogaster. — Malay. Nat. J. 16: 95—102.

Richards, O. W., 1962. A revisional study of the Masarid wasps. — Brit. Mus. (Nat. Hist.), 294 pp., 241
figs. — London.

———., 1971. The biology of the social wasps (Hymenoptera, Vespidae). — Biol. Rev. 46: 483—528.

———, 1972. The thoracic spiracles and some associated structures in the Hymenoptera and their sig-
nificance in classification, especially of the Aculeata. — Ent. Essays to commemorate reti-
rement Prof. K. Yasumatsu: I—13, 10 figs.

Saussure, H. de, 1852a. Description du genre /schnogaster. — Bull. Soc. ent. Fr. (2) 10: 19—27, 5 figs.

——— , 1852-8. Etudes sur la famille des Vespides, vols. 1 (1852-3), 2 (1853-8), 3 (1854—6). — Paris.

Smith, F., 1857. Catalogue of Hymenopterous Insects in the collection of the British Museum 5:
1—147. — London.

Spradbery, J. P., 1975. The biology of Stenogaster concinna van der Vecht with comments on the phylo-
geny of the Stenogastrinae (Hymenoptera: Vespidae). — J. Austr. ent. Soc. 14: 309—318, 12
figs.

Vecht, J. van der, 1972. A review of the new genus Anischnogaster in the Papuan region (Hymenoptera,
Vespidae). — Zool. Meded., Leiden 47: 240—256, 60 figs., | pl.

———, 1975. A review of the genus Stenogaster Guérin (Hymenoptera, Vespoidea). — J. Aust. ent.
Soc. 14: 283—308, 131 figs.

Williams, F. X., 1919. Philippine Wasp Studies II. Descriptions of new species and life history studies.
— Bull. Exp. Stn Hawaii. Sug. Plntrs Ass. (Ent. Ser.) 14: 19—186, 106 figs.

Yoshikawa, K., R. Ohgushi & S. F. Sakagami, 1969. Preliminary report on entomology of the Osaka
City University 5th scientific expedition to Southeast Asia 1966. With descriptions of two new
genera of stenogasterine wasps by J. van der Vecht. — Nature & Life in S. E. Asia 6: 153—182,
4 + 75 figs.

Der
RENE
hi

DEEL 120 AFLEVERING 3 1977

Di [568.2 MUS. COMP. zoo

VOOR ENTOMOLOGIE _

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

J. KRIKKEN. — Asian bolboceratine scarabs of the genus Bolbogonium Boucomont
(Coleoptera: Geotrupidae), p. 77—108, text-figs. 1—30, frontis-piece, pls. 2—3.

| Tijdschrift voor Entomologie, deel 120, afl.3 Gupubliceerd 1-VII-1977 |

Plate 1. Bolbogonium insidiosum, a new species from southern and central India (holotype male from
Coimbatore, total length ca 8.5 mm).

ASIAN BOLBOCERATINE SCARABS OF THE GENUS
BOLBOGONIUM BOUCOMONT (COLEOPTERA:
GEOTRUPIDAE)

by
J. KRIKKEN

Rijksmuseum van Natuurlijke Historie, Leiden

With 43 figures and one frontis-piece

ABSTRACT

After a brief introduction to and technical remarks on this first paper in a series on the classification
of Asian Bolboceratini, the genus Bolbogonium Boucomont is revised. Bolbogonium is here raised to
generic rank, diagnosed, and the 10 known species are keyed, described and illustrated. The
distribution of the genus is mainly Oriental. The three forms previously named are Bolbogonium
impressum (Wiedemann) comb. nov., punctatissimum (Westwood) comb. nov., and triangulum (West-
wood) comb. nov. (type-species). Of impressum and triangulum new records are given. The following
new species are treated: addendum (Vietnam), bicornutum (India), howdeni (India, Pakistan), insidiosum
(India), pseudopunctatissimum (India), scurra (India), and wiebesi (Burma).

A survey of characters and character states relevant to a supraspecific classification of the world
Bolboceratini is given in an appendix.

INTRODUCTION

This paper is the first in a series on the Asian representatives of the tribe
Bolboceratini (sensu Howden & Martinez, 1963). Both this series and related work
are intended to contribute to a world-wide reclassification of the group,
particularly on the supraspecific level.

During the past few years I have tried to assemble as many Asian bolboceratines
as possible, but the number of specimens so far received from many institutions
and individuals remained highly disproportional to my efforts, particularly from
the geographic point of view. This is, among other factors, certainly due to the
retiring habits of bolboceratines and to regional undercollecting; vast areas,
although in all probability inhabited by bolboceratines, do not seem to have
produced a single specimen. Consequently, the taxonomy and distribution of these
scarabs cannot be dealt with in a final or authorative manner, and sooner or later
supplementary results are to be expected.

Reliable data on the bionomics of the Asian species are not available, but

undoubtedly in the day-time the animals hide in deeply excavated terrestrial

|
|

burrows, as their European and North American relatives notoriously do (for a
summary of the known life-histories, see Howden, 1955: 296—299); they might as
well feed on and provide their brood burrows with subterranean fungi or other

Un

78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

vegetable matter. The label data indicate that several of the specimens before me,
were attracted to light; actually, the majority may have been collected in this way. |

The only comprehensive study on Asian Bolboceratini ever published was
written by Westwood (1852), who treated 22 species, omitting 7 names published
by earlier workers. Consultation of Westwood's specimens proved indispensable, |

but, unfortunately, part of the original material could not be traced. Later workers |
added 15 new species-group names based on material from tropical and eastern,

Asia, bringing the total to 44, of which two have been synonymized. The species |

were accomodated in seven genus-group taxa, including one subgenus. The last,

complete genus-group classification was given by Boucomont (1911) in a treatise

on the world fauna. He then proposed the subgenus Bolbogonium for Bolboceras |

triangulum Westwood. The taxonomy of this group, here raised to generic rank, is
worked out below.

The publication of a modern classification of the Asian genus-group taxa is
postponed, since this would entail the introduction of new names before the

taxonomy on the species-group level has been established to a reasonable degree.

A second reason for postponing is the fact that Howden and I are trying to work
out the classification of the world fauna. Some of the Asian groups are closely
related to or have members in the fauna of tropical Africa, and the taxonomic
characters of these are still under investigation. Despite this omission, the generic

diagnosis of Bolbogonium given hereafter warrants a correct recognition, as the .
features included implicitly match those of allied groups to be treated in

forthcoming papers.
In an appendix I give an analysis of the characters relevant to a comprehensive

classification of the world fauna, which may serve as a matrix for diagnoses of
genus-group taxa. Sixty-six characters are listed; 23 unconditional and 8 condi-

tional characters are suggested to be of primary importance in a supraspecific
classification. This does not necessarily mean that they are constant in all groups.
Although I have included classified character states derived from bolboceratines
of all the zoogeographic regions, the present survey is undoubtedly capable of
improvement and extension. The Australasian fauna seems incompletely surveyed,
at least as far as cephalic and pronotal armature is concerned. The classification of
the male genitalia is strongly simplified, mainly because of the lack of information
on the homologies of their elements. Most statements are phrased in a generalized
manner, and in actual cases need further specification. To give only one example,
I have no pretention at all to present a complete survey of the excessive diversity
in pronotal armature among Bolboceratini (characters 18 et seqq.). Only a few of
the characters listed can be used for phylogenetic research.

PRESENTATION OF DATA

The descriptive work was done with the aid of a Wild M5 binocular microscope
(magnifications x 6-x 100) plus drawing apparatus. Most distances and densities
were established by comparing the optical picture with verified scales viewed
through the drawing apparatus. Cephalic distances and densities were established
in full-face view, i.e. with the cephalic plane perpendicular to the optical axis.

KRIKKEN: Bolbogonium 79

Other measurements and counts, unless mentioned otherwise, were established
after placing the specimen with the plane touching the scutellum perpendicular to
the optical axis. The absolute values must be interpreted very carefully, since
measurements on such convex scarabs like Bolboceratini are inherently inexact.

The information in my descriptions varies slightly according to genus, number
and status of specimens available, and according to quantity and nature of
information in previous publications. In the present revision of Bolbogonium
detailed descriptions of all the species are given. As a rule little attention is paid to
the pectoral and abdominal characters. The abdomens of many specimens had to
be extracted in order to examine the genitalia. In Bolbogonium I suspect that at
least one species exhibits sexual dimorphism in its clypeofrontal ornamentation
(see below, section on the genus, infrageneric dissimilarities, character 2); in some
other species, however, the sexes are definitely similar. I noticed that some
workers had sexed their specimens incorrectly, this being due to the small size of
the phallus in the groups concerned; in several instances the abdomen had been
ruined completely, or it was simply missing.

Both the photographs of the general appearance and the drawings of the
cephalic and pronotal contours serve to relieve the descriptions of some
information difficult to communicate verbally. Unless mentioned otherwise, all the
scale lines given with the figures represent | mm. Some of the quantitative data in
the specimen descriptions have potential reference value only.

Clearly different size classes of sculptural elements (e.g. punctures) may be
mixed on a particular surface. In the descriptions this is indicated as double (two
size classes occur), triple (three size classes occur), etc., the elements being termed
primary (the largest), secondary, tertiary, etc. for each particular surface.

In Bolboceratini the number of fossorial elevations on the outer side of middle
and hind tibiae is difficult to establish. The height of these elevations decreases
from apex to base, and usually proximally only a number of spines indicate their
position. In the descriptions the number of indistinct proximal fossorial elevations
(usually those lacking a continuous crest) is placed between parentheses.

Despite poor collection data, the localities of the species are mapped; the
distribution of the genus (shaded, fig. 1) is hypothetical.

The following abbreviations concerning the location of Bolbogonium material
are used:

BH — Zoologisches Museum der Humboldt-Universität, Berlin;
BM — British Museum (Natural History), London;
CNC — Canadian National Collection, Ottawa;
— Rijksmuseum van Natuurlijke Historie, Leiden;
P — Muséum National d’Histoire Naturelle, Paris;
M — Zoologische Staatssammlung, Munich;
SMT — Staatliches Museum für Tierkunde, Dresden.

Genus Bolbogonium Boucomont stat. nov.

Bolbogonium Boucomont, 1911: 340 (as subgenus of Bolboceras Kirby; type-sp. Bolboceras triangulum
Westw.), 342 (in key).

80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

Generic diagnosis. — Middle coxae widely separated by anterior lobe of
metasternal disc, which is pyriform (fig. 28). Glabrous shiny area on proximal side
of club segment | large, and distinctly separated from surrounding pubescent
surface (fig. 27). Seven striae between suture and humeral umbone, all virtually
reaching base (fig. 26). Vertex laterally limited by arcuate crest, posterior margin
more or less concavely acclivous. Pronotum anteriorly with either simple or more
complex, characteristically shaped impression, usually surmounted by simple or
bisinuate crest (i.e., at most with single median protrusion).

Outer margin of right mandible with distinct arcuate lobe. Labrum thickened,
surface frequently heavily sculptured. Clypeal outline (dorsal view) variant:
semicircular, semielliptic or trapeziform, with or without anterolateral or antero-
median marginal protrusion(s). Perimarginal ridge of clypeus variably distinct.
Clypeus at most with low median longitudinal elevation; clypeofrontal transition
or frons with either single protrusion or set of protrusions, always situated discally.
Head never with simple transverse ridge between eyes. Frontovertex with or
without impression(s). Anterior border of eye-canthus marginate. Eyes entire, not

Fig. 1. Approximate known distribution of Bolbogonium (shaded) and its species. 1, addendum; 2,

bicornutum; 3, howdeni; 4, impressum; 5, insidiosum; 6, pseudopunctatissimum; 7, punctatissimum; 8,

scurra; 9, triangulum; 10, wiebesi. Numbers between parentheses refer to insufficiently detailed locality
data (usually country or province records).

KRIKKEN: Bolbogonium 81

divided into two parts. Pronotal base marginate (at least medially). Scutellum
virtually triangular, ratio 1/w 1—1.5. Elytral base unmodified; epipleuron reaching
apicosutural angle. Elytral interstriae scarcely convex, striae at most weakly
impressed. Antennal club unmodified; outline approximately ovate (ratio 1/w of
lamellae exceeding 1.5; fig. 27). Prosternum unmodified. Subapical fossorial
elevations of middle and hind tibiae (fig. 29) with either angulate or arcuate crest;
at least two complete non-apical fossorial elevations present. Metatarsus of middle
and hind legs relatively short, length not exceeding maximum width of tibial apex.
Parameres small, poorly sclerotized, simple (fig. 31). Body medium-sized, length
roughly around one cm. Colour uniformly brown, orange, or yellow.

Type-species. — Bolboceras triangulum Westwood, by original designation.

Affinities. — Bolbogonium species superficially resemble Bolboceras indicum
Westwood and its allies. They differ, however, greatly in characters of the
antennal club, striation on the elytral base, shape of metasternum, etc. The
primary set of properties, as mentioned in the first paragraph of the above
diagnosis, warrants a generic status, and Boucomont’s subgenus is here treated
accordingly.

Infrageneric dissimilarities. — The following properties proved to be important
in a classification of Bolbogonium (see comparative drawings):
1, shape of clypeus simply trapeziform (a), clypeus with produced anterolateral

angles (b), or clypeus more or less produced anteromedially (c).

2, detailed structure of clypeofrontal ornamentation; frons with variably de-
veloped transverse ridge (a), with two small, isolated (paramedian) tubercles
(b), with three small, isolated tubercles (c), with simple median tubercle (d),
with a different ornamentation (e), — note that this may be a sexual character.

3, frontolateral ridge distinct (a), or (sub)obsolete (b).

4, vertex generally flat (a), or with distinct U-shaped impression (b).

5, anteromedian impression of pronotum absent or superficial (a), or antero-
median impression well pronounced (b).

6, conditional on 5 (b); pronotal impression with large, well-defined (sub)hori-
zontal space immediately behind anterior border (a), or such a horizontal
space absent (b).

7, bisinuate crest of pronotum absent or ill-defined (a); or pronotum with distinct
crest, which is either moderately defined (b), or sharply defined (c).

8, basal margin of pronotum completely ridged (a), or incompletely ridged
(usually medially only) (b).

9, non-apical fossorial elevations on middle and hind tibiae with either arcuate
(a), or angulate (emarginate, bilobate) crest (b).

10, sculpture of dorsum — not classified.

The structure of the male genitalia is rather simple; they are generally poorly
sclerotized, and consequently provide no characters of practical diagnostic value.

The distribution of the classified character states over the species recognized
within Bolbogonium is given in the following table.

On these characters four species-groups may be recognized within Bolbogonium,
viz. the bicornutum group (with addendum and bicornutum), the triangulum group
(with pseudopunctatissimum, triangulum and impressum), the insidiosum group (with

=

82

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

Character no. l 2 3 4 5 6 7 8 9
addendum a a b a a | (b) a b a
bicornutum a a b a a (b) ab b a
pseudopunctatissimum b a a a a a a b a
triangulum b b/e a a b a a-b a a
impressum b d a a b a b a a
scurra a e ab a b b b b a
insidiosum c e a b b b b b a
punctatissimum c b b a b b b a b
howdeni c c b a b b b a b
wiebesi c d b a b b c a b

insidiosum and scurra), and the punctatissimum group (with punctatissimum, howdeni
and wiebesi).

There is no basis to discuss the phylogeny of Bolbogonium, since the significance |
of most characters is unknown.

Distribution. — Oriental, transgression into Palaearctic (fig. 1): 10 species

recorded from localities ranging from Pakistan to South Vietnam, no material seen
from North of the Himalaya.

considered casual.

w

Key to the species of Bolbogonium
(see figs. 2—25)

Anterolateral angles of clypeus distinctly raised, produced. Impression of

Bionomics. — Apparently nocturnally active; found “in soil”; record from dung ,

anterior side of pronotum with (sub)horizontal base. Vertex lacking sym- |
metrical impression(s). If there are only two frontal tubercles, these are not
connected by a conspicuous saddle. Subapical fossorial elevations of middle
and hind tibiae with arcuate crest. Pronotal base completely marginate |

(except in pseudopunctatissimum). Frontolateral ridge usually distinct. Eye-

canthus with more or less distinct anterolateralangle .............. p

Anterior border of clypeus either approximately straight with simply obtuse
anterolateral angles, or rounded with obsolete anterolateral angles. Im-

pression of anterior side of pronotum lacking well-defined horizontal base 5
Frons with well-pronounced median tubercle between eyes, and a longitudinal |
callosity on clypeus. Pronotal crest distinct, and shifted to posterior half of

pronotum. Length 9.5—13 mm. —N. India ......... impressum (p.92) |
Frons with set of twoorthree elevations)’. i. PP PRE 3%

the intergenal distance, connected by arcuate ridge. Pronotal crest ill defined,

. Frons between eye-canthi with pair of low tubercles separated by about half —

and shifted to posterior half of pronotum. Length 7.5—11 mm. — N. India.
LICH LIRA ee DEL COCA pseudopunctatissimum (p.90) :
Frons with different set of elevations, or with pair of approximated tubercles |

ee ee ee ee ee ee ee I ee © © © © © © © TO)

KRIKKEN: Bolbogonium 83

. Frons with pair of approximated tubercles. Length 7.5—10 mm. — Burma,
India»Rakistans et ee neem Bef triangulum 3 (p.91)
Frons with transverse ridge directly behind clypeofrontal suture, plus small
central tubercle. Length 7.5—12 mm. — Burma, India, Pakistan ........

ee Er eee triangulum © (p.91)
. Frons with pair of tubercles connected by variably developed rectilinear
elevation. Anterolateral angles of clypeus distinct. Pronotal base medially
feebly marginate. Frontolateral ridge indistinct ................. 6
Frons with 1—3 tubercles, either isolated or connected by more or less
pronounced V- or U-shaped elevation. Anterolateral angles of perimarginal
ridge of clypeus (sub)obsolete, and clypeal margin usually either rounded, or
with some anteromedian protrusion. If anterolateral angles distinct, frons with
V-shaped elevation. Elytral striae distinctly impressed ............. 7
. Frons with pair of widely separated small tubercles connected by vague ridge.
Pronotum slightly impressed behind anteromedian border. Posterior declivity
of vertex low. Length 8 mm. — S. Vietnam .......... addendum (p. 87)
Frons with pair of widely separated stout tubercles connected by saddle.
Anterior declivity of pronotum with characteristic impression. Length 11.5
MMS NES In dia rr A arene che A bicornutum (p. 89)
. Frons with single transverse tubercle between eye-canthi. W-shaped crest of
pronotum sharply defined. Clypeus with transverse antero-marginal costa.
Vertex lacking pronounced impression. Subapical fossorial elevations of
middle and hind tibiae with arcuate crest. Length 10 mm. — Burma ......

RARE di. Me Bern ee RE: wiebesi (p. 99)
Frons with two or three tubercles, free or connected by ridge ........ 8
. General elevation of clypeofrons with one anterior protrusion and a pair of
posterior tubercles; anterior protrusion frequently obsolescent, occasionally
only leaving its costiform connection between the posterior paramedian
tubercles; clypeofrontal suture (as far as visible) medially slightly shifted
foreward. Pronotal base medially feebly marginate. Frontolateral ridge usually
distinct. Subapical fossorial elevations on middle and hind tibiae with arcuate
erest&Vertcxösparsely.punctaten Anken OT 9
Frons with two or three small, isolated tubercles between eye-canthi. Pronotal
base completely marginate. Frontolateral ridge indistinct. Distal fossorial
elevations of middle and hind tibiae angulate-emarginate or bilobate. Vertex
denselyspunctate a. reo ene en N a 10
. Surface of vertex plane or nearly so. Elytral striae coarsely punctate,
punctures sharply defined, stria 2 obsolete slightly past scutellar apex. Clypeus
trapeziform, apex non-protuberant, slightly curved. Frontal elevation a wide-
legged'V. Length 8.5 —Ilmm. SC India .............. scurra (p. 94)
Surface of vertex symmetrically impressed, sparsely punctate. Elytral striae
moderately punctate, stria 2 extending further caudad. Clypeus with rounded,
more or less obsolete anterolateral angles, apex frequently protuberant.
Frontal elevation usually U-shaped, never a wide-legged V. Length 7—11 mm.
Sand. C: Indiages #4. 3a nation. ehe! insidiosum (p. 95)

84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

pseudopunctatissimum
impressum

bicornutum

triangulum Q

Bolbogonium addendum
triangulum Ô

Figs. 2—7. Contours of left half of head (dorsal), with approximate left side profile. 2, Bolbogonium
addendum, holotype; 3, bicornutum, holotype: 4, pseudopunctatissimum, holotype; 5, triangulum, &
Bengal; 6, ditto, @ Karachi; 7, impressum, 3 Bengal.

KRIKKEN: Bolbogonium 85

(©)
=

wiebesi

insidiosum

insidiosum 9
howdeni

scurra

punctatissimum

Figs. 8—15. Contours of left half of head (dorsal), with approximate left side profile. 8, Bolbogonium

scurra, holotype; 9, insidiosum, holotype; added figs. 10—12, polymorphism in shape of clypeofrons:

Nagpur (10), Hoshangabad (11), and Coimbatore (12); 13, punctatissimum, holotype; 14, howdeni,
holotype; 15, wiebesi, holotype.

86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

addendum |

pseudopunctati

18

bicornutum

17

punctatissimum

21

triangulum

20

impressum

19

insidiosum

scurra

22

Figs. 16—23. Contours of left half of pronotum (dorsal). 16, Bolbogonium addendum, holotype; 17,
bicornutum, holotype; 18, pseudopunctatissimum, holotype; 19, impressum, 3 Bengal; 20, triangulum, 3
Bengal; 21, punctatissimum, holotype; 22, scurra, holotype; 23, insidiosum, holotype.

KRIKKEN: Bolbogonium 87

. howdeni

wiebesi

a
a
»
3
>
se
3
è
8
a
$
Q
»
î
s
”
î

Figs. 24— 25. Contours of left half of pronotum (dorsal). 24, Bolbogonium howdeni, holotype; 25, wiebesi,

holotype. Figs. 26—31. Details of B. triangulum (26, 28—30, 3° Haldwani; 27, © Bengal; 31, ¢ Bengal).

26, left elytron, and scutellum; 27, flagellar segments and inward surface of first club segment; 28,

metasternal plate; 29, left hind leg; 30, left for leg; 31, phallus, dorsal. Scale line with fig. 27=0.5 mm,
others | mm.

10. Frons with three small transversely collinear tubercles between eye-canthi.
Head and pronotum crowdedly punctate or punctate-rugulate throughout.
Elytral derm moderately coarsely wrinkled, notably on lateral declivity.
Length 9-11 mm. — N. India, S. Pakistan ............ howdeni (p. 97)

— Frons with pair of small tubercles between eye-canthi. Pronotum densely or
crowdedly punctate. Lateral declivity of elytron not conspicuously wrinkled.
eneth, Sammy = Nain dias es re rg: punctatissimum (p. 96)

Bolbogonium addendum sp. nov. (figs. 2, 16, 32)

Description (holotype, female). — Approximate length 8, width 5, height 4 mm.
Orange-brown, shiny; tips, ridges, margins, sutures more or less infuscated;
pilosity yellowish. Habitus, fig. 32.

88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

Labrum short, almost rectilinear in front, sides rounded; surface rugulate.
Cephalic contours, fig. 2. Clypeus flat, marginal ridges distinct; surface punctate-
rugulate; clypeofrontal suture only laterally noticeable. Frons with pair of small,
widely separated tubercles connected by feeble, virtually rectilinear ridge; general

surface scarcely raised, between the eye-canthi irregularly crowdedly punctate,

almost punctate-rugulate; to the vertex this sculpture passes into double
punctation; primary punctures approximately isodiametric, rather coarse, well-

defined, closely and irregularly set, their diameters ca. 0.1 mm; secondary
punctures distinct, their diameters roughly one-fifth of those of the primaries, |

mostly separated by at least their own diameter; frontolateral ridge indistinct. Eye- |

canthus with raised anterior margin, sculpture punctate-rugulate. Maximum
length of head (exclusive of labrum) 1.95, maximum width 2.70 mm; ratio l/w 0.73.

Pronotal contours, fig. 16; surface of pronotum evenly convex, only surface,
immediately behind the anteromedian border depressed; anterolateral angles,
obtuse, posterolateral angles obsolete, widely rounded; pronotal base medially |

marginate. Pronotal punctation double; primary punctures approximately iso-
diametric, rather coarse, well defined and distinctly impressed, irregularly
distributed, closely set, except on paramedian parts of disc; densities on sublateral
surface ca. 30/sq. mm, their diameters ca. 0.1 mm; secondary punctures numerous,
distinct, evenly distributed. Median length of pronotum 2.8, maximum width 4.7
mm; ratio 1/w 0.60. Scutellum deltoid, with arcuate apex; surface virtually
impunctate.

Juxtasutural punctures of elytron small, indistinct. Discal striae of elytron

weakly impressed, striae on lateral declivity not impressed; punctures in striae

\

|

approximately isodiametric, distinct, densely arranged, separated by a few times

their diameters, which are ca. 0.05 mm; depressed peripunctural surface affecting
interstriae. Discal interstriae hardly convex, with numerous minute punctures.
Fore tibia with 7 external denticles, their size decreasing proximad; terminal
spur well developed, reaching approximately halfway tarsal segment 2. Femora all
moderately setose beneath, without notable details. Middle and hind tibiae with
spinose fossorial elevations increasingly developed distad; number on outer side of
right middle tibia 3 + (2), on right hind tibia also 3 + (2); crests of non-apical

elevations arcuate, apical one of middle tibia slightly emarginate; number of

fossorial spines along crest of anteapical elevation on right hind tibia ca. 17,

intermediate ones smaller than outer ones; spur of hind tibia scarcely tapering, |

nearly reaching tarsal segment 3.

Identification. — Bolbogonium addendum makes a comparatively pauperized

impression, and one wonders whether this has to do with marginal geographic
distribution or with mere individual variation. As with B. bicornutum, the dorsal

outline of the clypeus is trapeziform; the frontal elevation is diminutive compared |
to bicornutum. The posterior declivity of the vertex and the anterior impression of

the pronotum are ill pronounced.

Material examined. — Holotype only (P), which has the following label data: |
„Museum Paris/Cochinchine/Mont de Chaudoc/Harmand 1877”, ,,518/77”, ,,Mu-
seum Paris/Hte-Vera Paz/Bocourt 188—66”, „groupe de/Bolboceras triangulum ~

West./ Asie!// Bolboceras (Amechanus”. Paulian (1945: 41) already mentioned this

KRIKKEN: Bolbogonium 89

specimen. It undoubtedly belongs in Bolbogonium, and I think that the first-
mentioned label is the correct one.

Bolbogonium bicornutum sp. nov. (figs. 3, 17, 33)

Description (holotype, female). — Approximate length 11.5, width 6.5, height
5.5 mm. Orange-brown, shiny; tips, ridges, margins, sutures more or less
infuscated; pilosity yellowish. Habitus, fig. 33.

Labrum short, slightly emarginate in front, sides widely rounded, surface
rugulate. Cephalic contours, fig. 3. Clypeal margins raised; surface crowdedly
punctate to malleate-punctate; clypeofrontal suture distinct laterally. Frons with
pair of stout subconical tubercles between eye-canthi; tubercles connected by
transverse saddle steeply descending to clypeofrontal transition, posterior surface
gently sloping down to vertex; frons and vertex with distinct scattered, medium-
sized, isodiametric punctures; punctures crowded just below black tubercular tips,
elsewhere separated by a few times their diameters, which are ca. 0.05 mm;
density medially ca 10/0.25 sq. mm; between these punctures several smaller, ill-
defined, shallow punctures are noticeable; frontolateral ridge indistinct. Eye-
canthi with raised margin, surface punctate-rugulate. Maximum length of head
(exclusive of labrum) 2.55, maximum width 3.55 mm; ratio 1/w 0.72.

Pronotal contours, fig. 17; anterior declivity characteristically, shallowly
impressed; impression topped by gently declivous bisinuate crest, its median
prominence not sharply projecting cephalad, discal midline shallowly depressed;
anterolateral angles obtuse, posterolateral angles obsolete, widely rounded;
pronotal base medially marginate, laterally lined with punctures. Pronotal
punctation double; anterior impression, discal depression and lateral declivities
with numerous scattered, approximately isodiametric, well-defined, infuscated
punctures, mostly separated by at least one time their diameters, which are ca.
0.08 mm.; densities halfway lateral declivity ca. 20/sq. mm.; secondary punctures
sparse, moderately evenly distributed, their diameters diminutive compared to
primary ones. Median length of pronotum 3.9, maximum width 6.5 mm; ratio 1/w
0.59. Scutellum deltoid with slightly sinuate sides; surface with numerous small
scattered, distinct punctures, closely set in front, elsewhere sparse.

Elytron with only stria 1 shallowly impressed, others are merely series of
punctures; juxtasutural series of punctures well-developed. Strial punctures large,
resembling those of pronotum, regularly spaced, diameters on disc ca. 0.08 mm.,
mostly separated by two or three times their diameters. Interstriae 2 etc. not
noticeably convex, their surface with minute, indistinct, sparse punctures.

Fore tibia with 7 or 8 external denticles, their size decreasing proximad;
terminal spur of both fore tibiae missing. Femora all moderately densely setose
beneath, without notable details. Middle and hind tibiae with spinose fossorial
elevations increasingly developed distad; number in right middle tibia 3 + (3), in
right hind tibia 4 + (2); crest of non-apical elevations arcuate, apical crest of
middle tibia angulate; number of fossorial spines along crest of anteapical
elevation in right middle tibia 13; spines approximately equal-sized; only one
slightly bent, acuminate spur and two tarsal segments present in right middle tibia,
other accessories of middle and hind tibiae missing.

90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

Identification. — Bolbogonium bicornutum is easily recognizable by the shape of

the head, the dorsal outline of the clypeus being perfectly trapeziform, the frons |.

bearing a remarkably high transverse elevation. Furthermore, the outline of the
punctate impression on the anterior declivity of the pronotum is very character-

|

i
|
|
Il

in

istic. The elytral striation is most superficial, while the strial punctures are rather |

coarse, sharply defined.
Material examined. — Holotype only (M), from India: Calcutta.

Bolbogonium pseudopunctatissimum sp. nov. (figs. 4, 18, 34)

Description (holotype, male). — Approximate length 7.5, width 5, height 4 mm.

Brown, shiny; tips, ridges, margins, sutures more or less infuscated; pilosity |

yellowish. Habitus, fig. 34.
Labrum emarginate in front, sides widely rounded, surface rugulate-punctate.
Cephalic contours fig. 4. Clypeus with raised anterolateral angles; surface ru-

gulate-punctate; marginal ridge distinct; clypeofrontal suture distinct. Frons with |

pair of tubercles connected by low arcuate ridges; punctation of frontovertex |

double (x 25), contiguous in front, less dense on disc, where densities of primary

punctures are 13-17/0.1 sq.mm, diameters ca. 0.08 mm; punctures well defined, |

|
|
\
|

distinctly impressed, isodiametric; frontolateral ridge distinct. Eye-canthus with
weakly raised anterior margin, surface contiguously punctate. Maximum length of |

head (exclusive of labrum) 1.90, maximum width 2.40 mm; ratio 1/w 0.79.
Pronotal contours, fig. 18; anterior declivity only with subhorizontal base,

lacking impression; transverse discal crest subobsolete, discal midline shallowly _ |

ti
|

|

impressed; base medially marginate. Pronotal punctation double, laterally triple
(x 50); primary punctures sparse on paramedian discal surface; secondary

punctures large, isodiametric, distinctly impressed, tertiary punctures well defined,
isodiametric, equal-sized; densities of primary punctures sublaterally 8-12/0.25 sq.
mm, diameters of primary, secondary and tertiary punctures 0.15, 0.08 and 0.01
mm respectively. Median length of pronotum 2.5, maximum width 4.6 mm; ratio
1/w 0.54. Scutellum deltoid, moderately punctate.

Juxtasutural stria of elytron present; discal striae of elytron shallowly impressed.
Strial punctures distinctly impressed, well defined, isodiametric, their diameters
ca. 0.05 mm, separated by 1-2 times their own diameter; peripunctural impressions
affect interstrial surface. Interstria 1 distinctly convex, other discal interstriae very
weakly convex; punctation sparse, secondary punctures just visible at magnifi-
cation x 50.

Fore tibia with 7 external denticles, their size decreasing proximad; terminal
spur well developed, reaching to apex of tarsal segment 2. Femora all moderately
densely setose beneath, without notable details. Middle and hind tibiae with
spinose fossorial elevations increasingly developed distad; number in right middle
tibia 3 + (2), in right hind tibia 3 + (2); crest of non-apical elevations arcuate,
apical elevation of middle tibia emarginate; number of fossorial spines along crest
of anteapical elevation on right middle tibia ca. 10, all nearly equal-sized; spurs of
hind tibia slightly tapering but with rounded apex, reaching to apex of tarsal
segment 2.

KRIKKEN: Bolbogonium 9]

Variation. — Length 7.5-11 mm. Holotype with obsolescent pronotal crest,
which is slightly better defined in the paratypes.

Identification. — Because of its bituberculate frons, this species was confound-
ed with Bolbogonium punctatissimum. It differs from that species by its produced
clypeal angles, the ridge connecting the frontal tubercles, the poorly pronounced
pronotal crest and the arcuate fossorial elevations on middle and hind tibiae. The
frontal and pronotal characters mentioned also separate pseudopunctatissimum
from its closest relative, triangulum. Contrary to the other species in the triangulum
group, the pronotal base of pseudopunctatissimum is incompletely marginate.

Material examined. — 3 specimens.

Holotype with label reading ,,Musoorie/Mackenzie coll./10.vii-20.x.22” (BM).
Paratypes, 13° from Himalaya (BM), and 14 from Himachal Pradesh: Simla, viii-
1898 (SMT).

Bolbogonium triangulum (Westwood) comb. nov.
(figs. 5, 6, 20, 26-31, 35-37)

Bolboceras triangulum Westwood, 1852: 26 (type-loc. Mussoree), pl. 4 figs. 20, 20a.
Bolboceras (Bolbogonium) triangulum; Boucomont, 1911: 340 (type-sp. of the subgenus); Boucomont,
1912: 14 (in catalogue).

Notes. — There are two groups of specimens here placed under triangulum,
which at first sight differ by their frontal ornamentation (figs. 5, 6). One group,
agreeing with the type, proved to consist of females; the other proved to consist of
males; sometimes both forms were found in the same series. As no other
morphological differences could be found, I consider the frontal ornamention in
this species a sexual character, despite the fact that such dimorphism was not
found in other species. Furthermore, there is some variation in the arrangement of
the frontal protrusions (compare figs. 35 and 36). More material is needed to
confirm the present interpretation of B. triangulum.

Description. — Approximate length 3, 7.5-10, 9, 7.5-12 mm. Brown, orange or
yellow; shiny; pilosity yellowish. Habitus, figs. 35-37.

Labrum short, emarginate in front, sides widely rounded, surface rugulate-
punctate. Cephalic contours, figs. 5(¢), 6 (2). Clypeal surface rugulate-punctate;
anterolateral angles of clypeus produced; clypeofrontal suture distinct. Frons with
pair of approximated tubercles (Z), or with transverse, at the ends tuberculate
ridge plus median interocular tubercle (9); frequently with elongate callosity on
clypeofrontal transition; frontolateral ridge distinct, extending to acute crest
limiting vertex; anterior surface of frons rugulate-punctate; posterior surface
punctate, punctures usually more abundant in Z. Eye-canthus rugulate-punctate,
with raised anterior margin.

Pronotal contours, fig. 20; anterior declivity with well-defined horizontal base,
topped by distinct W-shaped crest plus lateral cavity; midline distinctly impressed,
well defined behind median protrusion; pronotal borders entirely marginate.
Pronotal punctation triple; secondary punctures very distinct, their diameters ca.
one-fifth of the primaries; tertiary punctation scarcely distinct (x 50); primary
punctation dense along discal midline, very dense or crowded laterally, punctures

92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

deep, well defined; anterior declivity microreticulate, opaque. Scutellum (fig. 26)

deltoid, densely, distinctly punctate.

Elytral contours, fig. 26; juxtasutural punctures fine. Elytral striae discally |

weakly impressed; punctures moderately defined, separated by 1-3 (or 4) times

their diameters. Interstriae weakly convex, with very fine punctures, separated by ©

several times their diameters.

Fore tibia (fig. 30) with 7 external denticles, their size decreasing proximad; —

terminal spur reaching to tarsal segment 2. Femora all moderately setose beneath,
without notable details. Middle and hind tibiae (fig. 29) with spinose fossorial

elevations increasingly developed distad, two or three non-apical elevations having |

a complete arcuate crest with ca. 15 approximately equal spines; terminal spurs
well developed, approximately as long as tarsal segments | + 2.

Identification. — Bolbogonium triangulum females are immediately recognizable
by their characteristic frontal ornamentation. Males (as interpreted here, see
above) may be confounded with other species with a bituberculate frons. Within
the triangulum group with its produced anterolateral clypeal angles, only pseudo-
punctatissimum has two frontal tubercles, but these are less approximated than
in triangulum, and they are connected by a transverse ridge. In triangulum the
pronotal crest is nearly always more strongly pronounced than in pseudopunctatis- |
simum. The secondary punctures on the pronotum of triangulum are much larger in
proportion to the primaries than in pseudopunctatissimum.

Material examined. — 36 specimens.

Holotype from India with label „Ind. or./Mussoure/in cow dung” (BM). I doubt
if this ecological remark indicates a regular habit. Further specimens as follows (if
sex is indicated this was established by extraction of genitalia).

India: Bengal, no further details (2 ¢, SMT, one labelled by Boucomont as B.
punctatissimum); Bara Taunda [?], x. 1927, Konietzko (1, BH); Buxar (2 &, 1 9,
SMT); Cawnpore [Kanpur], 12.1.1921, Vernon (1, BM); Chapra, Mackenzie (2 9,
BM); Deccan (1, P); Dehra Dun, 13.vii.1928 (14, BM); Dhara, v.1943 (1 9,
Forest Research Inst. and Colleges, Dehra Dun); Calcutta (1, M); Haldwani:
Chakata Range, 23.vi.1930, Chatterjee (1 ¢, L); Kumaon (1 ©, BM); Motinala R.,
25.vi.1927), Chatterjee (1 9, BM); Naldera, 29.vi.1938, Beeson, in soil (1 9, Forest
Research Inst. and Colleges, Dehra Dun); Pusa, 5.xii.1904. Watson (1 9, BM),
17.vii.1915, at light (1 9, BM), 1.1.1916, Bahadur, at light (1 ©, BM), 2.iii.1920,
Austin (1 9, BM), Pusa without further data (1 ©, BM); Simla (1 ©, BM). North
India, no details (1 &, BM). — Burma, (1 g, seen by Boucomont, | © with
perfectly straight frontal ridge, BM). — Pakistan: Karachi, Bell (3 ¢, 1 9, 1, BM);
Murree (1 3, seen by Boucomont, BM); Quetta, vii.viii.1933, Samuel (1 ©, BM),
vii.1936, Nazeer, peach plot, light trap (1 9, BM; frontal elevations poorly
pronounced); Tarnab, 24.v.1916, Fletcher (1 &, BM; fig. 35).

Bolbogonium impressum (Wiedemann) comb. nov. (figs. 7, 19, 38)

Scarabaeus impressus Wiedemann, 1823: 6 (type-loc. Bengal).
Bolboceras impressum; Boucomont, 1902: 5 (in catalogue); 1912: 10 (in catalogue).

KRIKKEN: Bolbogonium 93

Description. — Approximate length 9.5-13 mm. Brown, largely shiny; pilosity
yellowish. Habitus, fig. 38.

Labrum short, slightly emarginate in front, sides widely rounded, surface
rugulate. Cephalic contours, fig. 7. Clypeus shallowly concave behind strongly
produced anterolateral angles; midline with elongate callosity interrupting dis-
tinct, virtually rectilinear clypeofrontal suture; clypeal surface entirely malleate-
rugulate. Frons raised between eye-canthi, medially with stout tubercle, the apex
of which may be slightly bifid; surface malleate-rugulate, posteriorly normally
punctate; frontolateral ridge distinct. Vertex shallowly depressed, arcuate lateral
margins distinctly raised; punctation of vertex double; primary punctures
scattered, approximately isodiametric, shallow but distinct, laterally closely set,
medially mostly separated by a few times their diameters; secondary punctures
minute, scarcely discernable (x 50), but numerous, evenly distributed. Antero-
lateral angle of eye-canthus raised, sculpture like that of clypeus and frons.

Pronotal contours, fig. 19; anterior side with conspicuous, more or less opaque
impression; declivity topped by virtually W-shaped crest with distinct median
protrusion; base of declivity virtually horizontal, its posterior limit nearly
parallelling superior crest; surface in front of lateral part of superior crest with
rugulate-punctate cavity; pronotal borders entirely marginate. Pronotal punc-
tation triple; primary punctation sparse beside impressed discal midline, close to
crowded laterally, anteriorly, and on discal midline; punctures distinct, mostly
isodiametric, laterally more or less irregular; secondary punctures very small, their
diameters about one-tenth of the primaries, evenly distributed discally; tertiaries
scarcely noticeable (x 50). Scutellum deltoid; closely punctate, secondary
punctures minute, numerous, evenly distributed.

Juxtasutural punctures of elytron fine. Discal striae of pronotum shallowly
impressed; punctures infuscated, small, separated by 2-4 times their diameters.
Interstriae very weakly convex, punctation double; primary punctures sparse,
scattered, small, approximately isodiametric, separated by several times their
diameters; secondary punctures minute, numerous, evenly distributed.

Fore tibia with 8 external denticles, their size decreasing proximad; terminal
spur reaching to tarsal segment 2. Femora all moderately setose beneath, without
notable details. Middle and hind tibiae with spinose fossorial elevations increasing-
ly developed distad, two or three non-apical ones having a complete, arcuate
crest; anteapical crest with 15-20 approximately equal spines; terminal spurs well
developed, nearly as long as tarsal segments | + 2.

Identification. — Bolbogonium impressum, the largest species in the genus, has a
single well-pronounced transverse tubercle on the middle of the frons, by which it
is immediately separated from the other species in the triangulum group. B. wiebesi
bears a similar frontal tubercle, but differs in shape of clypeus, sharpness of
pronotal crest, general sculpture of dorsum, etc.

Material examined. — 6 specimens.

Holotype from Bengal (Copenhagen museum). Further specimens as follows.

India: Dehra Dun, leg. Asmadon (1); Pusa, i.1921, leg. Senior White 145),
16.1x.1916, leg. Fletcher (1g); Simla, vii.1909 (19) (all BM); another specimen
from Bengal without details (14, SMT), figured here.

94 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

Bolbogonium scurra sp. nov. (figs. 8, 22, 39)

Description (holotype, male). — Approximate length 11, width 6, height 5 mm.
Orange-yellow, shiny; tips, ridges, margins, sutures, punctures more or less
infuscated; pilosity yellowish. Habitus, fig. 39.

Labrum long, emarginate in front, sides widely rounded, surface rugulate-
punctate. Cephalic contours, fig. 8. Clypeus with distinctly raised anterior margin,

lateral ridges distinct; surface irregularly rugulate-punctate. Frontal elevation ©

situated immediately against clypeofrontal suture, which is slightly shifted
foreward; elevation consisting of a V-shaped saddle connecting a pair of
infuscated tips; anterior declivity of elevation very steep, closely, finely punctate,
posterior declivity gently sloping to vertex, punctate-rugulate; lateral declivity
with similar sculpture. Disc of frontovertex virtually flat, very sparsely punctate,

sides and posterior declivity of vertex more densely punctate; punctures

approximately isodiametric, moderately defined, distinctly impressed, their dia-

meters ca. 0.05 mm, densities centrally scarcely exceeding 5/0.1 sq. mm. Lateral |
delimitation of vertex formed by distinct, but not particularly raised, arcuate crest. —

|
|
|

i

Eye-canthus with raised anterior margin, sculpture punctate-rugulate, adjacent |
section of frontolateral ridge obsolescent. Maximum length of head (exclusive of.

labrum) 2.50, maximum width 3.15 mm; ratio 1/w 0.80.
Pronotal contours, fig. 22; anterior declivity of pronotum impressed, but lacking

distinct horizontal base behind apical border; superior crest moderately pro-
nounced, median protrusion distinct, discal midline shallowly impressed; antero- |

lateral angles obtuse, posterolateral angles obsolete, widely rounded; base
marginate. Pronotal punctation double; base of anterior declivity abundantly |
punctate, sides closely punctate; remaining surface also with abundant primary

punctures, except around protrusion and on paramedian parts of disc, where
secondary punctation is dominant; primary punctures large, isodiametric, distinct-
ly impressed and generally well-defined; diameters of sublateral punctures ca. 0.07
mm (increasing laterad), their densities 7-10/0.25 sq. mm (increasing laterad);
secondary punctures fine, isodiametric, sizes variant. Pronotal median length 3.1,
maximum width 6.1 mm; ratio 1/w 0.50. Scutellum deltoid, finely punctate;
micropunctures (x 75) present.

Juxtasutural punctures of elytron very distinct. Discal striae of elytron shallowly
impressed; punctures infuscated, deeply impressed, exceedingly well-defined,
isodiametric, locally irregularly spaced, their diameters ca. 0.05 mm; peripunctural
impressions indistinctly affecting interstrial surface; stria 2 abbreviated behind.
Interstriae very weakly convex, with numerous fine punctures, generally separated
by a few times their diameters, which are ca. 0.07 mm.

Fore tibia with 7 external denticles, their size decreasing proximad; terminal
spur well developed, reaching halfway tarsal segment 2. Femora all setose
beneath, without notable details. Middle and hind tibiae with spinose fossorial
elevations increasingly developed distad; number in right middle tibia 3 + (2), in
right hind tibia also 3 + (2); crests of non-apical elevations arcuate; number of
fossorial spines along crest of anteapical elevation on right middle tibia 12, all

KRIKKEN: Bolbogonium 95

approximately equal-sized; spurs of hind tibiae with rounded tip, not tapering,
reaching segment 3 of tarsus.

Variation. — Length 8.5-11 mm. The three specimens at hand are, apart from
the usual differences related to size, very similar.

Identification. — Bolbogonium scurra is close to insidiosum, from which it differs
primarily by its flat vertex, its different V-shaped frontal elevation (the legs of the
V being wide apart), and its trapeziform clypeus.

Material examined. — 3 specimens.

Holotype from South India: Coimbatore, vi.1966, leg. P.S. Nathan (Howden
collection). Two paratypes, also from Coimbatore, xi.1966, leg. P.S. Nathan
(Howden collection, L).

Bolbogonium insidiosum sp. nov. (figs. 9—12, 23, 40)

Description (holotype, male). — Approximate length 8.5, width 5, height 3.5
mm. Yellow-brown, shiny; tips, ridges, margins, sutures, punctures more or less
infuscated; pilosity yellowish. Habitus, plate 1.

Labrum emarginate in front, sides widely rounded, surface rugulate-punctate.
Cephalic contours, fig. 9. Clypeus surface rugulate-punctate; sides limited by
ridge, protuberant anteromedially. Frontal elevation situated inmediately against
clypeofrontal suture, which is slightly shifted foreward; elevation U-shaped,
anteriorly protuberant, on either side terminating at well-developed tubercle;
interior surface of U punctate-rugulate, elevated; sides of frons punctate-rugulate
as well; frontolateral ridge distinct. Vertex with large U-shaped, sparsely punctate
impression, laterally limited by acute crest. Eye-canthus with raised anterior
margin, surface indistinctly rugulate. Maximum length of head (exclusive of
labrum) 2.10, maximum width 2.50 mm; ratio 1/w 0.85.

Pronotal contours, fig. 22; anterior declivity impressed, but without horizontal
base behind apical border; superior crest moderately pronounced, median
protrusion distinct, discal midline shallowly impressed; anterolateral angles
obtuse, posterolateral angles obsolete, widely rounded; base marginate medially.
Pronotal punctation generally sparse, double (x 25), more abundant on midline
and lateral declivities; primary punctures large, isodiametric, distinctly impressed,
generally well defined; their densities sublaterally 10-12/sq. mm, diameters
somewhat less than 0.1 mm. Median length of pronotum 2.45, maximum width
4.75 mm; ratio 1/w 0.52. Scutellum deltoid, finely punctate; micropunctures (x 75)
present.

Juxtasutural punctures of elytron distinct. Discal striae of elytron shallowly
impressed; punctures large, deeply impressed, well defined, locally irregularly
spaced, their diameters ca. 0.05 mm, separated by 2—4 times this diameter;
peripunctural impressions slightly affecting interstrial surface; stria 2 extending
onto posterior declivity. Discal interstriae weakly convex with fine punctures,
generally separated by few times their diameter.

Fore tibia with 8—9 external denticles, their size decreasing proximad; terminal
spur well developed, reaching halfway tarsal segment 2. Femora all setose
beneath, without notable details. Middle and hind tibia with spinose fossorial

96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

elevations, increasingly, developed distad; number in right middle tibia 2 + (3), in |
right hind tibia 3 + (2); crests of non-apical elevations arcuate; number of |

fossorial spines along crest of anteapical elevation on right middle tibia ca. 15;

Spurs of hind tibiae with rounded tip, not tapering, reaching tarsal segment 3.
Variation. — Length 7—11 mm. This species is extremely variable in shape of

clypeus, ornamentation of frons and vertex, development of the anteromedian

|

protrusion of pronotum. The variation in the shape of the clypeofrons is illustrated |

in figs. 1O—12.

Identification. — Bolbogonium insidiosum is recognizable by its frontal elevation |

and the characteristically impressed vertex. As stated in the preceding paragraph,

variation is considerable.

The frons may be trituberculate with variably pronounced intervening ridges.
The development of the tubercles themselves varies as well, the anterior one
independently from the posteriors; in extreme cases only an arcuate ridge is left in
front of the posterior pair of tubercles. The degree of impression of the vertex

|

varies strongly, but remains noticeable, at least in the specimens I have seen. _
Compared to scurra, two useful features are the greater extension of elytral stria 2

and the frontal elevation not being a wide-legged V.

Material examined. — 33 specimens.

Holotype from India: Madras: Coimbatore, xi.1964, leg. P.S. Nathan, 1400 ft
(M). Paratypes as follows.

India. — Bellary, 1896, De Morgan (1, P, mentioned by Boucomont, 1911: 340);
Belgaum (1 9, BM); Chinchona, Anomalai Hills, v.1966, Nathan, 3500 ft (1 9,
Schulze coll.); Coimbatore, xi.1955 (4, CNC), xi.1958 (1, Howden coll.), xi.1962 (1,
CNC), x.1962 (2, CNC), xi.1963 (1 9, L), i.1964 (6, CNC), xii.1966 (6, Howden
coll.), all Nathan, 1400 ft; Hoshangabad, 14-19.ix.1911, T.S.F. [?], at light (1 9,
BM); Kadanpur [?, indistinct handwriting] (1 9, 1, SMT); Kadegaon (1 &, 1 9,
BM, seen by Boucomont); Nagpur, 10.xii.1915 (1 9), 2.xii.1917 (1), 26.xii.1918 (1),
d’Abreu (all BM); Buldana, Sagoda Purna, 7.iii.1930, Chatterjee (1, BM). A worn
© from ,,India bor.” (BM) excluded from type-series (mentioned by Boucomont,
1911: 341).

Bolbogonium punctatissimum (Westwood) comb. nov. (figs. 13, 21, 41)

Bolboceras punctatissimus Westwood, 1852: 22, pl. 4 fig. 9, 9a (type-loc. Moradabad).
Bolboceras punctatissimum; Boucomont, 1912: 12 (in catalogue).

Description (holotype, not sexed). — Approximate length 8, width 5, height 4
mm. Brownish yellow, shiny; tips, ridges, margins, sutures more or less infuscated;
pilosity yellowish. Habitus, fig. 41.

Labrum short, slightly emarginate in front, sides widely rounded, surface
sculpture indistinct. Cephalic contours, fig. 13. Clypeal margins raised, particular-
ly anteriorly, marginal ridge obsolete near rounded anterolateral angles; surface
closely punctate, punctures small, somewhat irregular, shallow, weakly defined;
interspaces not exceeding diameters of punctures, which are ca. 0.05 mm; clypeo-
frontal suture laterally distinct. Middle of frons raised between genal angles, with

KRIKKEN: Bolbogonium 97

pair of low transverse elevations; punctation like that of clypeus, but generally
closer, almost malleate-punctate; frontolateral ridge indistinct. Lateral delimit-
ation of vertex not marginate, though still abruptly declivous; posterior surface of
vertex concavely acclivous; sculpture like that of clypeus and frons. Eye-canthus
with raised margin, sculpture like that of frons. Maximum length of head
(exclusive of labrum) 1.8, maximum width 2.4 mm; ratio 1/w 0.79.

Pronotal contours, fig. 21; anterior declivity impressed; superior crest poorly
pronounced, discal midline shallowly depressed, anterolateral angles obtuse,
posterolateral angles obsolete, widely rounded; borders entirely marginate.
Pronotal punctation double (x 50); base of anterior declivity and sides punctate-
rugulate, remaining surface with close primary punctation; punctures small,
shallow, locally ill-defined; diameters of discal punctures mostly slightly exceeding
0.05 mm, their densities 50-60/0.25 sq. mm; secondary punctures numerous,
extremely fine. Median length 2.8, maximum width 4.7 mm; ratio 1/w 0.58.
Scutellum deltoid, its surface punctate.

Juxtasutural punctures of elytron indistinct. Discal striae shallowly impressed;
punctures slightly infuscated, small, shallow, regularly spaced, peripunctural
impressions indistinctly affecting interstrial surface. Interstriae scarcely convex,
with scattered primary punctures mostly separated by one or two times their
diameters, which are ca. 0.025 mm; secondary punctation as on pronotum.

Fore tibia with 7 external denticles, their size decreasing proximad; terminal
spur well developed, reaching to tarsal segment 3. Femora all moderately setose
beneath, without notable details. Middle and hind tibiae with spinose fossorial
elevations increasingly developed distad; number in right middle tibia 3 + (2) in
right tibia 3 + (2); crests of distal elevations emarginate; number of fossorial
spines along crest of anteapical elevation in right middle tibia 6 (superiorly) and
ca. 9 (inferiorly), inferior spines longer than superior ones; spurs of middle tibia
acuminate; those of hind tibia scarcely tapering, with rounded tip, nearly as long
as tarsal segments I + 2.

Identification. — Bolbogonium punctatissimum is closely allied to B. howdeni but
differs in frontal ornamentation and general sculpture of dorsum. These dif-
ferences are rather tentative, since the type of punctatissimum is the only specimen
known to me. Some forms in the triangulum group have also a pair of small frontal
tubercles, but these are readily distinguished by the produced anterolateral angles
of the clypeus.

Material examined. — Holotype only (BM), with label reading “Ind. or./
Moradabad/on Evening/in the Ruins/by a Candle”.

Bolbogonium howdeni sp. nov. (figs. 14, 24, 42)

Description (holotype, male). — Approximate length 9, width 5, height 4 mm.
Light brown, shiny; tips, ridges, margins, sutures more or less infuscated; pilosity
yellowish. Habitus, fig. 42.

Labrum rather strongly protruding, almost rectilinear in front, sides rounded;
surface rugulate-punctate. Cephalic contours, fig. 14. Clypeus shallowly concave
behind raised front margin, lateral borders not distinctly raised; surface entirely

98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

malleate-punctate; clypeofrontal suture indistinct. Raised clypeofrontal disc with
3 small but conspicuous tubercles between genal angles; frontal sculpture similar
to that of clypeus; frontolateral ridge indistinct. Vertex closely punctate, punctures
approximately isodiametric, small (diameters ca. 0.05 mm), shallow, but distinct.
Eye-canthus with raised anterior margin, sculpture rugulate. Maximum length of
head (exclusive of !abrum) 1.80, maximum width 2.40 mm; ratio 1/w 0.76.

Pronotal contours, fig. 24; anteromedian declivity topped by distinct bisinuate .
crest; only surface immediately behind anteromedian border distinctly depressed; |

discal median longitudinal depression shallow; anterolateral angles obtuse,
posterolateral angles obsolete, widely rounded; pronotal borders entirely mar-

ginate. Pronotal disc crowdedly punctate, punctures approximately isodiametric, |

well defined, of variable sizes, their diameters rarely reaching 0.1 mm; lateral
declivities punctate-rugulate. Median length of pronotum 2.8, maximum width 5.1
mm; ratio 1/w 0.57. Scutellum deltoid, apex shortly arcuate; surface closely
punctate.

Juxtasutural punctures of elytron weakly impressed. Striae on lateral declivity of
elytron indistinct owing to strongly transversely wrinkled surface; punctures of
striae 2 et seqq. poorly defined, regularly spaced, their diameters ca. 0.1 mm
mostly separated by less than three times this diameter. Discal interstriae scarcely
convex, with scattered small but distinct punctures, their diameters ca. 0.05 mm;
micropunctation hardly visible ( x 50).

Fore tibiae with 7 external denticles; terminal spur well developed, reaching
halfway segment 2. Femora all moderately setose beneath, without notable details.
Middle and hind tibiae with spinose fossorial elevations increasingly developed
distad; number in right middle tibia 3 + (1), in right hind tibia 3 + (2); crest of
distal elevations emarginate; number of fossorial spines along crest of anteapical
elevation ca. 4 (superiorly) and ca. 4 (inferiorly), their sizes variant; spurs of hind
tibiae not tapering, with rounded tip, reaching approximately halfway tarsal
segment 2.

Variation. — Length 9-11 mm. The four specimens at hand are, apart from the
usual differences related to size, very similar.

Identification. — Bolbogonium howdeni is easily recognizable by the presence of
three small tubercles on the frons and by its heavily sculptured dorsum
(contiguously punctate, malleate-punctate, or punctate-rugulate). B. punctatis-
simum is certainly closely allied with howdeni, but clearly differs in the aforesaid
characters.

Material examined. — 4 specimens.

Holotype from India: Bihar; Pachrukhi, 1927 (Howden collection); Howden
correctly labelled it as being allied with punctatissimum. Paratypes from India:
Chapra, leg. Mackenzie (1 9, BM); Pusa, 30.xi.1904 (1, BM). — Pakistan: Sind (1
©, P).

Note. — This species is dedicated to Dr. H. F. Howden, specialist in
Geotrupidae, professor of biology at Carleton University, Ottawa.

KRIKKEN: Bolbogonium 99
Bolbogonium wiebesi sp. nov. (figs. 15, 25, 43)

Description (holotype, male). — Approximate length 10, width 6.5, height 4.5
mm. Brown, shiny; tips, ridges, margins, sutures more or less infuscated; pilosity
brownish. Habitus, fig. 43.

Labrum quite prominent, slightly emarginate in front, sides widely rounded,
surface indistinctly rugulate. Cephalic contours, fig. 15. Front margin of clypeus
strongly raised, forming a transverse costa; lateral limiting crests of horizontal
surface obsolete. Frons with very weakly bifid transverse tubercle; clypeofrontal
suture vaguely distinct laterally; frontolateral ridge indistinct. Eye-canthus with
raised outer margin. Cephalic surface almost entirely malleate-punctate, im-
pressions on posterior declivity of frontal tubercle somewhat transversely
confluent; separate units medium-sized, their diameters just behind the eyes
slightly over 0.05 mm. Maximum length of head (exclusive of mouthparts) 2.45,
maximum width 2.40 mm; ratio 1/w 0.83.

Pronotal contours, fig. 25; anterior side with distinct impression topped by W-
shaped crest; discal midline very shallowly impressed; anterolateral angles obtuse,
posterolateral angles obsolete, widely rounded; borders entirely marginate.
Pronotal punctation triple (magnification x 50); disc and anterior declivity with
close primary punctation, punctures rather coarse, shallowly impressed though
well defined, approx. isodiameteric; their diameters ca. 0.07 mm, densities just
beside midline ca. 10/0.25 sq. mm; density of primary punctures increasing
laterad, marginal zone punctate-rugulate; secondary punctures discally most
distinct, scattered, approximately isodiametric, diameters ca. 0.03 mm, densities
beside discal midline almost 20/0.25 sq. mm; tertiary punctures numerous,
moderately evenly distributed on pronotal disc. Median length of pronotum 3.4,
maximum width 6.2 mm; ratio 1/w 0.55. Scutellum deltoid, with scarcely sinuate
sides; surface with numerous distinct punctures, slightly smaller than primary ones
of pronotal disc, diameters ca. 0.05 mm; secondary punctation dense, punctures
resembling tertiaries of pronotal disc.

Juxtasutural punctures of elytron indistinct. Striae discally lightly impressed;
punctures small, diameters scarcely exceeding 0.05 mm, separated by a few
diameters; peripunctural impressions slightly affecting interstrial surface. Inter-
striae very slightly convex, punctation double; primary punctures scattered,
distinct, approximately isodiametric, diameters less than 0.05 mm, separated by
at least one diameter; secondary punctation remarkably close, punctures evenly
distributed, fine, diameters less than one-tenth of the primary ones.

Fore tibia with 7 external denticles, their size decreasing proximad; terminal
spur of fore tibia well developed, extending a little beyond tarsal segment 1.
Femora all setose beneath, without notable details. Middle and hind tibiae with
spinose fossorial elevations increasingly developed distad; number in right middle
tibia 3 + (2); distal crests emarginate, number of fossorial spines along crest of
anteapical elevation 5 (+ ca. 5 long setae) (superiorly), and 5 (inferiorly); spurs of
middle tibia acuminate, those of hind tibia scarcely tapering, with rounded tip,
slightly shorter than tarsal segments | + 2.

Identification. — Bolbogonium wiebesi is recognizable by the large frontal

100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

tubercle; impressum, similar in this character, differs primarily by the produced
anterolateral angles of its clypeus. In wiebesi the anterior margin of the clypeus is |
strongly costate; furthermore, the pronotal crest is remarkably sharp and the, |
entire dorsum is heavily punctate. |

Material examined. — Holotype only (BM), with labels reading ‘‘Burma/1919- |
103.515 |

Note. — This species is dedicated to Dr J. T. Wiebes, professor of systematic |
zoology and evolutionary biology at the State University of Leiden.

NOTES ADDED IN PROOF

Two recent descriptions of Bolbogonium extend the generic range (fig. 1) some
hundreds of kilometers into Afghanistan. Bolbogonium kabakovi Nikolajev (1976: «
693) from the Laghman province confirms the introductory notes given above
under B. triangulum Westw., and I suspect a synonymy here. Bolbogonium
kabulicum Nikolajev & Kabakov (1977: 646) from Kabul would easily key to point
7 in my key, and then seems immediately recognizable by its peculiar frontal
ornamentation (l.c.: fig. 1).

ACKNOWLEDGEMENTS

My protracted study of Asian Bolboceratini is possible only thanks to the
patient assistance of several colleagues and their technicians. Relevant to the
material dealt with in this paper are:

M. E. Bacchus (London), A. Bons (Paris), P. N. Chatterjee (Dehra Dun), A.
Descarpentries (Paris), H. Freude (Munich), F. Hieke (Berlin), H. F. Howden —
(Ottawa), R. Krause (Dresden), N. Mgller Andersen (Copenhagen), R. D. Pope |
(London), J. Schulze (Berlin). |

Critical comments on drafts of this paper were received from H. F. Howden |
(Ottawa), P. J. Kuijten (Leiden), J. T. Wiebes (Leiden).

The habitus drawing (plate 1) was produced by A. Bos, the photographs were
printed by C. Hoorn, both on our museum staff.

Visits to institutions abroad were made possible by grants from the Uytten-
boogaart-Eliasen Stichting (Amsterdam).

REFERENCES

Boucomont, A., 1902. Coleoptera Lamellicornia, Fam. Geotrupidae. — Genera Insectorum 7, 20 pp. |
pl.

Boucomont, A., 1911. Contribution a la classification des Geotrypidae (Col.). — Annls Soc. ent. Fran-
ce 79 (1910): 335—350.

Boucomont, A., 1912. Scarabaeidae: Taurocerastinae, Geotrupinae. — Col. Catalogus 46, 47 pp.

Howden, H. F., 1955. Biology and taxonomy of North American beetles of the subfamily Geotrupinae,
with revisions of the genera Bolbocerosoma, Eucanthus, Geotrupes, and Peltotrupes (Scarabaei-
dae). — Proc. U.S. nat. Mus. 104: 151—319, pls. 1—18.

Howden, H. F. & A. Martinez, 1963. The new tribe Athyreini and its included genera (Coleoptera: Sca-
rabaeidae Geotrupinae). — Canad. Ent. 95: 345—352, figs. 1—19.

KRIKKEN: Bolbogonium 101

Nikolajev, G. V., 1976. Neue Arten der Blatthornkäfer (Coleoptera, Scarabaeidae) aus Afghanistan. —
Polskie Pismo ent. 46: 693—697, figs. 1—3.

Nikolajev, G. V. & O. N. Kabakov, 1977. Four new species of the family Scarabaeidae (Coleoptera)
from Afghanistan. — Zool. Zhurnal 56: 646—647, figs. 1—5. (In Russian).

Paulian, R., 1945. Coléoptères Scarabéides de l’Indochine. 1. — Faune de l’Empire français 3, 228 pp.,
figs. 1—105, 1 map.

Westwood, J. O., 1852. Descriptions of some new or imperfectly known species of Bolboceras, Kirby. —
Trans. linn. Soc. London 21: 19—30, pls. 3—4.

Wiedemann, C. R. W., 1823. Zweihundert neue Kafer von Java, Bengalen und den Vorgebirgen der
Guten Hoffnung. — Zool. Mag. 2: 1—135.

102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

APPENDIX

CHARACTERS AND CHARACTER STATES RELEVANT TO A SUPRASPECIFIC |
CLASSIFICATION OF THE TRIBE BOLBOCERATINI !)

Head (dorsal)

1 (a) Outline of mandibles (outer margins) symmetrical.
(b) Outline of mandibles (outer margins) asymmetrical.
2 (a) Outer margin of right mandible sinuate, poorly lobed, or simply arcuate.
(b) Outer margin of right mandible, for the greater part, straight, or nearly so
(c) Outer margin of right mandible with distinct arcuate lobe.
(d) Outer margin of right mandible with distinct acute lobe.
3 (a) Transverse ridge of labrum present and distinct.
(b) Transverse ridge of labrum absent, obsolete, or at least indistinct.
*4 (a) Outline of clypeus in dorsal view (male) approximately trapeziform.
(b) Outline of clypeus in dorsal view (male) approximately semicircular. |
(c) Outline of clypeus different from preceding alternatives, e.g., very strongly modified by | |
marginal protrusion(s) — to be specified for each case.
*5 (a) Perimarginal ridge of clypeus present and unmodified.
(b) Perimarginal ridge of clypeus present and modified — but different from alternative (d). |
(c) Perimarginal ridge of clypeus entirely or largely obsolete. |
(d) Perimarginal ridge transformed into X-shaped ridges (dorsal view), cross-point with or |
without tubercle. |
*6 (a) Base of any cephalic protrusion(s) restricted to clypeus, frons, or vertex, or situated on:
clypeofrontal transition.
(b) Base of usually well-developed cephalic protrusion largely covering both clypeus and frons.
7 (a) Clypeus lacking single anteromarginal protrusion.
(b) Clypeus with single anteromarginal protrusion — but not as sub character state 4 (c).
*8 (a) Clypeal disc with transverse ridge or pair of isolated elevations.
(b) Clypeofrontal transition (suture) with transverse ridge or pair of isolated elevations.
(c) Clypeal disc with a single tubercle — but not as sub character state 6 (b).
(d) Clypeofrontal transition (suture) with a single tubercle — but not as sub character state 6 (b).
*Q (a) Frontal disc with transverse ridge or pair of isolated elevations.
(b) Frons with pair of distinct para-ocular protrusions.
(c) Frons or vertex with a single tubercle.
10 (a) Frontovertex lacking distinct impression(s), at most shallowly concave.
(b) Frontovertex with distinct impression(s).
*11 (a) Vertex lacking transverse ridge.
(b) Vertex with distinct transverse, straight or arcuate ridge.
*12 (a) Vertex not sharply separated from tempora.
(b) Vertex separated from tempora by ridge. |
*13 (a) Tempora not produced.
(b) Tempora more or less produced, angulate.
14 (a) Vertex gently sloping to pronotal apex.
(b) Vertex posteriorly limited by vertical declivity. |
15 (a) Anterior margin of eye-canthus simply arcuate. |
(b) Anterior margin of eye-canthus straight, shortly rounded or with lateral angle and/or tubercle.
*16 (a) Eye-canthus and temporal lobe separated.
(b) Eye-canthus and temporal lobe contiguous, dividing eye in two parts.
17 (a) Dorsally visible area of eye small — to be specified for each case.
(b) Dorsally visible area of eye large — to be specified for each case.

1) Characters suggested to be of primary diagnostic importance are marked with an asterisk.

KRIKKEN: Bolbogonium 103
|Pronotum (male) and scutellum

18 (a) Pronotum lacking distinct non-marginal protrusions, simply convex (except for shallow median
longitudinal sulcus).

(b) Pronotum with distinct non-marginal protrusions — characters 19—22 conditional on this
| character state.
Fa (a) Pronotum simply retuse anteromedially, declivity surmounted by transverse discal crest;
| lacking discal paramedian or lateral protrusions.

(b) Pronotum distinctly concave anteromedially, but lacking transverse discal crest or any other

protrusions.
(c) Pronotum with pairs of paramedian and/or lateral protrusions (usually tubercles); in well-
developed forms frequently with anteromedian and/or sublateral impressions.

we (a) Pronotum lacking crest just behind apex proceeding onto lateral declivities.

(b) Pronotum with crest just behind apex proceeding onto lateral declivities.
*21 (a) Pronotum not, at the same time, with sharp posterior crest running roughly parallel to base,

and anterior surface retuse to strongly concave.

| (b) Pronotum with sharp posterior crest running roughly parallel to base, and anterior surface
| retuse to strongly concave.
| 22 (a) Pronotal protrusion(s) not restricted to basal area.
| (b) Pronotal protrusion(s) restricted to basal area.
|
|
|

1*23 (a) Pronotal apex lacking pair of small deep impressions.
(b) Pronotal apex with pair of small deep impressions (bifoveate).
| 24 (a) Pronotal base marginate (at least medially).
(b) Pronotal base immarginate.
1*25 (a) Pronotal apex medially marginate or immarginate, but unmodified.
| (b) Pronotal apex modified (e.g., tuberculate or with high transverse ridge).
| 26 Conditional on 25 (b).
(a) Pronotal apex unituberculate (in one known case with long upbent horn).
(b) Pronotal apex bituberculate, occasionally tubercles confluent or obsolescent.
(c) Pronotal apex strongly elevated over much of its width (costate or carinate).
27 (a) Pronotal apex fringed with narrow velum and/or setae.
(b) Pronotum lacking velum, but fringed with long setae.
1*28 (a) Scutellum semicircular, semielliptic, sides may be more or less parallel or weakly sinuate in
front — character state 29(c) exepted.
(b) Scutellar sides strongly sinuate (curving laterad in front) — character state 29(c) excepted.
(c) Scutellum simply triangular.
(d) Shape of scutellum different form preceding alternatives — to be specified for each case.
*29 (a) Scutellum very wide (1/w<1).
(b) Scutellum short (1/w 1—2).
(c) Scutellum elongate (1/w>2).

Elytron

*30 (a) Elytral base immarginate.
(b) Elytral base marginate (ridged).
31 (a) Elytral epipleuron reaching apico-sutural angle.
(b) Elytral epipleuron obsolete at some distance from elytral apex (at the beginning of the distal
curve).
*32 (a) Elytron with 7 striae between suture and humeral umbone.
(b) Elytron with 5 striae between suture and humeral umbone.
(c) Elytron with 9 striae between suture and humeral umbone.
33 (a) Stria 5 extending to (near) elytral base.
(b) Stria 5 obsolete in front.
*34 (a) Stria 1 terminating at side of scutellum, stria 2 interrupted in front or reaching base of elytron
(not basal angle of scutellum).
(b) Both striae 1 and 2 terminating at side of scutellum (2 occasionally terminating at basal angle
of scutellum).

104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

(c) Stria 1 proceeding to elytral base. |
35 (a) Elytral striae 2—7 not or only scarcely impressed, intervals (= interstriae) not or only scarcely)
convex. |
(b) Elytral striae sulcate, intervals distinctly convex.
36 Conditional on 35(b). |
(a) Elytral intervals all approximately equally convex. |
(b) Odd intervals of elytral disc (at least juxtasutural one) much more convex than even intervals. | |
37 (a) Humeral angle of elytron unmodified.
(b) Humeral angle of elytron tuberculate.

Antenna

38 (a) Antennal club normal, not thicker than length of pedicel and subsequent flagellar segments |
combined. |
(b) Antennal club very robust, thicker than length of pedicel and subsequent flagellar segments | I
combined (outer surface of ultimate segment usually distinctly convex). |
*39 Conditional on 38 (b). |
(a) Distal side of club segment 3 evenly convex or nearly so. I

(b) Distal side of club segment 3 remarkably swollen near base.
*40 (a) Proximal surface of club segment | entirely pubescent, lacking glabrous, polished area.
(b) Glabrous, polished area on proximal side of club gradually passing to pubescent parts. |
(c) Glabrous, polished area on proximal side of well separated from remaining, pubescent surface. |
*41 (a) Distal side of club segment 3 lacking grooves. |
(b) Distal side of club segment 3 with 2 distinct grooves. |

Pectus

42 (a) Anterior paramedian costae of prosternum distinct.
(b) Anterior paramedian costae of prosternum obsolete.
43 Conditional on 42 (a).
(a) Prosternum convex or flat (occasionally wrinkled) between anterior paramedian costae.
(b) Prosternum with median longitudinal ridge, but juxtacoxal surface not deeply concave.
(c) Prosternum, behind anterior paramedian costae, with longitudinal costa limited by deep,
juxtacoxal holes. |
(d) Prosternum sulcate (not simply concave) between anterior paramedian costae.
44 (a) Prosternum lacking any posteromedian protrusion, not even a longitudinal ridge. |
(b) Prosternum with some posteromedian protrusion.
45 Conditional on 44 (b). |
(a) Prosternum lacking posteromedian spine or similar protrusion.
(b) Prosternum angulate, or with isolated posteromedian spine or similar protrusion.
*46 Conditional in 44 (b).
(a) Prosternum not bulbose or otherwise inflated-dilated.
(b) Prosternum bulbose posteromedially. |
(c) Prosternum with well-developed lanceolate posteromedian process. |
(d) Prosternum with transverse, bidentate posteromedian process.
*47 (a) Middle coxae (sub)contiguous, anteromedian process of metasternum indistinct. |
(b) Middle coxae distinctly separated, although anteromedian process of metasternum may be
strongly narrowed.
48 Conditional on 47(b).
(a) Metasternal process between middle coxae not strongly narrowed. |
(b) Metasternal process between middle coxae strongly narrowed.
*49 Conditional on 47(b).
(a) Metasternal disc pyriform in outline.
(b) Metasternal disc rhomboid in outline.
(c) Metasternal disc different from preceding alternatives — to be specified for each case.

KRIKKEN: Bolbogonium 105

*50 Conditional on 47(b).
(a) Metasternal process flat, or nearly so, abruptly retuse in front, with complete perimarginal ridge,
adjacent mesosternal declivity more or less concave.
(b) Metasternal process flat, or nearly so, abruptly retuse in front, lacking perimarginal ridge,
adjacent mesosternal declivity more or less concave.
(c) Metasternal process long and narrow (linear), but not reduced to a sharp carina.
(d) Metasternal process gradually passing to mesosternum, not limited by some transverse
| protrusion, lacking longitudinal carina.
(e) Metasternal process with distinct median longitudinal carina, which may be more or less
| angulate in front (prow-shaped).
(f) General surface of metasternal process hunched, different from preceding alternatives.
*51 Conditional on 47(b).
(a) Metasternal process lacking any particular accessory protrusions.
| (b) Metasternal process with robust denticle in front.
| (c) Metasternal process with fine spine or hook in front.
|

52 (a) Mesometasternal suture noticeable at base of declivity.
} (b) Mesometasternal suture obsolete.
53 (a) Longitudinal suture of metasternum (largely) obsolete.
(b) Longitudinal suture of metasternum distinct.

‚Abdomen

| 54 Structure of stridulatory organ — available data insufficient for classification.

Legs (male)

| 55 (a) Fossorial elevations of middle and hind tibiae with straight, angulate — emarginate, bidentate,
of bilobate crest (reference: anteapical elevation of hind tibia).
| (b) Fossorial elevations of middle and hind tibiae with arcuate crest (reference: ditto).
| 56 (a) Middle tibia with a single non-apical fossorial elevation, others completely obsolete or
noticeable only as pairs of isolated denticles.
(b) Middle tibia with two or more complete non-apical fossorial elevations.
| 57 (a) Fore tibia normal.
(b) Fore tibia strongly dilated.
1*58 (a) Femora indentate.
(b) Femora dentate (particularly fore femora).
\*59 (a) Terminal spur of fore tibia long and slender, scarcely tapering.
(b) Terminal spur of fore tibia more or less triangular, with acute apex, occasionally very robust.
*60 (a) Tarsal segment 1 of fore-legs short.
(b) Tarsal segment 1 of fore-legs remarkably elongate.

Genitalia

461 (a) Aedeagus consisting of two parameres.
(b) Separate parameres indistinct.
| 62 Conditional on 61 (a).
| (a) Parameres strongly sclerotized, without notable accessory elements.
(b) Parameres sclerotized and strongly modified, i.e. with accessory elements.
(c) Parameres very small, poorly sclerotized, without notable accessory elements.
(d) Different from preceding alternatives — to be specified for each case.

| 63 (a) Armature (sculptural ornamentation) of head and pronotum of male and female belonging to
the same species similar.
(b) Armature (sculptural ornamentation) of head and pronotum of male and female belonging to

the same species radically different — to be specified, see characters 4 et seqq., 18 et seqq.

|
|
|
|
Is exual dimorphism
|
|

106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977 |
Colours |
64 (a) Colour uniform throughout, apart from infuscated ridges, sutures, tips, margins, punctures, |
etc. |
(b) Colour not uriform thoughout.
65 Conditional on 64(a).
(a) Colour some lighter tone of brown.
(b) Colour some very dark tone of brown, or black.
66 Conditional on 64(b).
(a) Elytra and pronotum differently, but uniformly coloured.
(b) Elytra and/or pronotum with some colour pattern.

KRIKKEN: Bolbogonium 107

Plate 2. Bolbogonium species. 32, addendum, holotype; 33, bicornutum, holotype; 34, pseudopunctatis-
simum, holotype; 35—37, triangulum, 3 Tarnab (35), d Burma (36), © Kanpur (37), length ca 8, 11.5,
11 mm, respectively.

108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 3, 1977

Plate 3. Bolbogonium species. 38, impressum, & Bengal, length 12.5 mm; 39, scurra, holotype; 40,
insidiosum, holotype; 41, punctatissimum, holotype; 42, howdeni, holotype; 43, wiebesi, holotype.

RN
NEN

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Pal Din
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DEEL 120 AFLEVERINGEN 4 EN 5 1977

TIJDSCHRIFT una,
VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

F. WILLEM»E. — A study on the genus Cranaella Ramme (Orthoptera, Acridoidea,
Catantopinea), p. 109—120, text-figs. 1—30, pl. 1. — A study on the genus
Cranae Stal (Orthoptera, Acridoidea, Catantopinae), p. 121—152, text-figs.
1—45, pls. 1—5.

Tijdschrift voor Entomologie, deel 120, afl. 4 en 5 Gepubliceerd 1-VII-1977

A STUDY ON THE GENUS CRANAELLA RAMME
(ORTHOPTERA, ACRIDOIDEA, CATANTOPINAE)

by

FER WILLEMSE
Laurastraat 67, Eygelshoven, The Netherlands

With 30 text-figures and one plate

ABSTRACT

Morphological characters of all previously recognized species of the genus Cranaella Ramme are
given, together with the descriptions of three new species: kevani, rammei, and samarensis. A key is
given to the species of this genus, which occurs in the Philippines and Celebes.

INTRODUCTION

Material of Cranaella is scarce. Ramme, when creating this genus, recorded four
adult females and one juvenile male, representing three new species: carnipes,
tuberculata and willemsei. C. Willemse (1956) mentioned another two females and
two males, one male as willemsei, the other specimens as tuberculata. Kevan (1966)
recorded another two males and one female as tuberculata and described a fourth
species, multicolor, after a single female. Before me I have a collection of 30 males
and 17 females. In the present study morphological characters, especially of the
genitalia, are given, and three new species are described.

Depositories ofthe material are given throughout the text in abbreviated form:
ANSP Academy of Natural Sciences of Philadelphia, U.S.A.;

BPBM Bernice P. Bishop Museum, Honolulu, Hawaii;
NMM Natuurhistorisch Museum, Maastricht, Netherlands;
ZMHU Zoologisches Museum der Humboldt-Universität, Berlin, Germany.

My thanks are due to the following persons: J. L. Gressitt and the late Miss S.
Nakata, Honolulu; K. K. Günther, Berlin; H. Radclyffe Roberts and D. Rentz,
Philadelphia.

Cranaella Ramme
Cranaella Ramme, 1941: 94, 217, 226, 241; C. Willemse, 1956: 9, 108.
Type-species: Cranaella carnipes Ramme, 1941.

Cranaella belongs to the Cranae group of genera, preliminarily arranged under
the subfamily Catantopinae. In the original description it is said that Cranaella

109

110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 4, 1977 |
differs from Cranae Stal in the rougher sculpturation of head and thorax, the less |
strongly developed pronotal sulci and narrower elytra, with the hind margin (not |
“Vorderrand”) slightly concave. However, as to these points the discontinuity
between these genera is merely gradual. Also the genitalia are much the same in |
both genera. It is not within the scope of the present study to discuss the status of,
the genera of the Cranae group. Therefore I abstain from redescribing Cranaella |
more definitely. However, phenetic relationships of Cranaella species both
mutually as well as with species of the Cranae complex (s.l.) are pointed out here. !

In Cranaella, specific characters may be found in the apex of the phallus and the |
hind margin of the last abdominal tergite in the male, in the subgenital plate in the |
female and in the sculpturation of the hind femur, in the shape of the prosternal |
process, in the development of the hind wing and the shape of the elytron, and i in, |
the coloration in both sexes. |

Key to the species of Cranaella

1. Hind femur sanguineous (Celebes) ............ carnipes Ramme (p. 112) |
— Hind femurnotred (Philippine Is). SASSI 2 | |
2. Elytron wider, about two-and-one-half times as long as wide, yellowish green |
without black venation, reaching the margins .... multicolor Kevan(p. 119) |
— Elytron narrower, three times or more as long as wide, venation by far not |
reaching the (slightly transparent) margins, the area of venation black ... 3,
3. Ridges of the fishbone pattern of the outer medial area of the hind femur of |
same colour as the hind femur; each ridge with an, often black, tubercle near |
the lower and usually also near the upper carinulae; prosternal process |
widened laterally, lateral edge of apex slightly conical; hind wing vestigial . . 4 |
— These ridges in black, contrasting with the colour of the hind femur and
without tubercles; prosternal process almost cubical, lateral edges of the apex |
not conical; hind wing present, although shorterthantheelytron........ sh
4. Hind margin of last abdominal tergite in male not or slightly incised in the
middle, without furculae (fig. 20); apex of phallus with distal part distinctly |
marked off from the proximal part of the apex (figs. 10—11) ............
Ne BR ER ot Le TEE tuberculata Ramme (p. 112) |
— This margin with a pair of small and widely separated furculae (fig. 21); apex of |
phallus with distal part gradually merging in the proximal part (figs. 12—13) . . :
ee irk iors ee iio any cae EI AE O E e Sara kevani sp.n.(p. 115)
5 Hind margin of the last abdominal tergite in male shallowly emarginate in the
middle, without furculae (fig. 22); apex of phallus with short distal part (figs.

VEE De EEE eh willemsei Ramme (p. 116)
— This margin with a narrow incision in the middle and a pair of furculae (figs.
23—24); distal part of apex of phallus longer (figs. 16—19) ........... 6

6. Distal part of apex of phallus comparatively long (figs. 16—17) ..........
SOC he Che NK on 6 rammei sp.n.(p. 118)
— Distal part of apex of phallus comparatively short (figs. 18—19) .........
aia meh hese cy Leene VE NE MERE IE gh Wah sel a Meth co hee ee samarensis sp.n.(p. 118)

F. WILLEMSE: The genus Cranaella 111

Figs. 1—5. Cranaella tuberculata Ramme, g, Surigao, phallic complex: 1, dorsal view, ectophallic

membrane and epiphallus removed; 2, the same, lateral view: 3, endophallus, lateral view; 4,

epiphallus, dorsal view; 5, the same, posterior view. Figs. 6—9. Cranaella willemsei Ramme, g, Surigao,

phallic complex: 6, lateral view; 7, the same, epiphallus and ectophallic membrane removed, dorsal
view; 8, the same, lateral view; 9, endophallus, lateral view.

2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 4, 1977

Cranaella carnipes Ramme, 1941
(fig. 25)

Cranaella carnipes Ramme, 1941: 94, 217, 228, pl. 14 fig. 3; C. Willemse, 1956: 108, 109.

This species is known from the type-specimens, an adult female (holotype) and a
juvenile male (paratype). Both are labelled: Celebes Ile Ile 500 m 11.12.1930 G.

Heinrich (ZMHU). Ramme’s description is extremely short; his description of the |

coloration refers to the juvenile male; that of the holotype was given by C.
Willemse.

Morphological characters of the holotype are as follows: integument finely

pitted, impressions closely set; prosternal process widened laterally, lateral edges
slightly conical; elytron about three times as long as wide, hind wing vestigial;
ridges of outer medial area of hind femur not tuberculate; subgenital plate (fig. 25)
flattened ventrally and depressed apically, hind margin tridentate with the lateral
points almost as wide and long as the median point.

The juvenile male has the hind margin of the last abdominal tergite with a pair

of small, triangular and widely separated furculae, resembling those of kevani. The
tegmina are not yet developed. The phallic complex is so poorly sclerotized that |

details cannot be given.

Although the prosternal process and the tegmina are much as in tuberculata and |

kevani, the relationship of carnipes remains an open question by lack of an adult
male.

Cranaella tuberculata Ramme, 1941
(figs. 1—5, 10—11, 20, 26, pl. 1 figs. 1—2)

Cranaella tuberculata Ramme, 1941: 95, 217, pl. 13 fig. 6; C. Willemse, 1956: 108, 109.

Material studied: Surigao, Mindanao, Baker 4¢ 39; Surigao, P.I., 9.vi (19) &
9.v.1916 (1); Surigao, Mindanao 2¢ 1 juv. g, 29 1 juv.@; Iligan, Mindanao,
Baker 14; Butuan, Mindanao, Baker | g 1 2; Bucas, Philipp. 1 &; Siargao,
Philipp. 1 9; Island Samar, Baker 24 19; Isl. Biliran, Philippines, Baker 1g;
Dagami, Leyte, P.I., Mt. Lobi 21 (13) & 25 (19) & 26.vi(19) & 423) & 10.viii
(1 juv. 3) & 26 (13) & 27.ix.1945 (19), E. R. Helwig (all ANSP, except 14 29
from Surigao in NMM).

This species was described after the female holotype from Mindanao and a
female paratype from Siargao. A description of the male characters was given by
C. Willemse after a male from Surigao (Mindanao). The redescription which
follows here is made after (topotypic) material from Mindanao.

Redescription.

3, pl. 1 fig. 1. Integument of face, cheeks, thorax and dorsal sides of proximal
abdominal tergites evenly and strongly pitted. Head as long as the pronotum.
Pronotum with four transverse sulci, the fourth one located at about 5/6 of the
length of the middle of the dorsum; posterior margin of dorsum broadly
emarginate; lateral lobe 1/4 longer than high, posterior angle slightly produced

F. WILLEMSE: The genus Cranaella 113

Plate 1, figs. I—7. Cranaella species: 1, tuberculata Ramme, 4, Surigao; 2, idem, ©, Surigao; 3, kevani
sp.n., & holotype; 4, willemsei Ramme, &, Dinagat; 5, rammei sp.n., & paratype; 6, idem, 9 paratype;
7, samarensis sp.n., & holotype.

114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 4, 1977

posteriorly, ventral and posterior margins sigmoid. Prosternal process strong, |
vertical, slightly widened laterally, anterior side slightly longer than posterior one, |
ventral side flattened with the anterior margin straight or slightly convex, the |
posterior margin strongly convex, being curved towards the anterior margin, the |
lateral edges slightly produced conically. Elytron narrow, three to three-and-one- |
half times as long as wide, venation strongly reduced and confined to the middle ‘|
area; the broad margins almost completely transparent, anterior margin slightly ‘|
convex, posterior margin slightly concave, both margins about parallel, apex |
broadly rounded. Hind wing vestigial. Tympanum open, almost circular, on level |
with the body surface. Outer side of hind femur with the carinulae slightly |
tuberculate and the lower and upper ends of the ridges of the fishbone pattern |
more conspicuously tuberculate. Brunner’s organ present. Lower inner carinula |
pilose. |
Hind margin of the last abdominal tergite (fig. 20) widely concave, slightly |
thickened near the middle and shortly and narrowly incised in the very middle. |
Supra-anal plate (fig. 20) wider than long. Phallic complex and epiphallus as in
figs. 1—5. Apex of phallus (figs. 10—11) with the ventro-posterior side strongly
concave and the distal part of the apical penis valve distinctly marked off from the
proximal part of the apex by a lateral collar-shaped fold.

General colour yellowish or reddish-brown. Antennae bluish-black, apical |
segments slightly paler, basal segments coloured as the head. Head of general |
colour or olivaceous green, clypeus with a median black dot and its lateral angles ,
black. Thorax of general colour except for the ventral side of the prosternal |
process, the posterior margin of the pro- and the lateral margins of the |
mesosterna, which are bluish. Elytron with the area of venation black or dark blue.
Abdomen of general colour, last tergite, cercus and margins of the supra-anal |
plate dark brown or blue. Fore and middle legs and hind femur of general colour |
or more reddish. Hind femur with the tubercles of the ridges of the fishbone
pattern black; the lower inner marginal area bluish; a yellow or greenish |
antegenicular ring, often bordered with black proximally. Hind knee bluish-black,
apex of the lobes yellowish and crescents castaneous-brown. Hind tibia and its
condylus bluish-black, with a yellow postgenicular ring; spines with tips black.
Hind tarsus bluish-brown, ventral side paler.

©, pl. 1 fig. 2. Larger than male. Ventral side of subgenital plate depressed in the
middle, the depression bordered laterally by a pair of strong and posteriorly
slightly divergent keels, giving rise to a tridentate hind margin with the lateral
points smaller than the median point (fig. 26). Coloration as the male.

Measurements (length in mm): body, & 21.1-25.2, 9 23.5-32.0; pronotum, ¢
3.8-4.3, Q 4.9-5.9; elytron, & 4.1-4.5, © 4.9-6.0; hind femur, g 11.1-12.1, 9 13.0-
95%

Distribution: Philippine Is. (Mindanao; Bucas; Siargao; Samar; Leyte; Biliran).

Discussion. The species is well-defined by the male genitalia. Among the
material at hand, morphological characters vary but slightly. However, as to the
coloration, the material from Samar and that from Leyte and Biliran is different
from that of Mindanao. The pronotum of the Samar specimens has two black dots,

F. WILLEMSE: The genus Cranaella 115

at the lateral ends of the second transverse sulcus. The material from Leyte and
Biliran shows more or less numerous black dots scattered over the head and
pronotum and almost unicolorous tubercles of the outer medial area of the hind
femur.

Cranaella kevani sp.n.
(figs. 12—13, 21, 27, pl. 1 fig. 3)

Cranaella tuberculata (nec Ramme); Kevan, 1966: 411.

Material studied: ¢ holotype, 9 allotype, labelled: P.I., Misamis Or., Mt.

11 13 15 17 19

Figs. 10—19. Cranaella species, 3, apex of phallus in ventro-posterior (odd numbers) and lateral (even
numbers) view: 10—11, tuberculata Ramme, Surigao; 12—13, kevani sp.n., Calian; 14—15, willemsei
Ramme, Surigao; 16—17, rammei sp.n., paratype; 18—19, samarensis sp.n., holotype.

116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 4, 1977

Empagatao, 1050—1200 m, 19-30.iv.61, H. Torrevillas collector; paratypes: P.I.,
Misamis Or., Minalwang, 1050 m, 24.iii-4.iv.1961, W. Torrevillas 18 (all three
specimens BPBM and labelled: Cranaella tuberculata Rm det.D.K.McE.Kevan,
1965); Calian, Davao Prov., Mindanao, P.I., 10.v.1930, C. F. Clegg 1g; Island of,
Basilan, Baker 24 19 (ANSP). |

Description. |
&, pl. 1 fig. 3. Differs from tuberculata as follows. Pronotum slightly shorter, |
posterior margin of dorsum slightly emarginate. Hind margin of last abdominal |
tergite emarginate in the middle, with a pair of short and well separated furculae
(fig. 21). Distal part of the apical penis valve not strongly marked off but more
gradually merging in the proximal part of that valve (figs. 12—13). |
Coloration as in tuberculata. |
©. Larger than male. Subgenital plate resembling that of tuberculata, but the
keels slightly closer together (fig. 27). Coloration as in male, slightly mo
obscurely brownish. |
Measurements (length in mm): body, & 21.1-24.8, 9 27.0-30.9; pronotum, |
& 4.0-4.1, 9 5.0-5.3; elytron, G 3.8-4.6, 9 4.5-5.5; hind femur, & 11.6-13.0, |
© 13.9-15.0. |
Distribution: Philippine Is. (Mindanao; Basilan).
Discussion. The species is well-defined by the male genitalia. The furculae may |
be slightly more widely separated than in the holotype. Specimens from Basilan |
differ slightly in the smaller measurements, the presence of a black dot at either |
end of the second transverse pronotal sulcus, and the unicolorous elytron in the |
female. |
The material from Misamis Or. had been referred to tuberculata by Kevan. In |
that paper, the locality of the firstly recorded male (now paratype) is erroneously | |
given similar to that of the following couple (now holo- and allotype). |
Within the genus, tuberculata and kevani form a group characterized by the | |
shape of the prosternal process, vestigial hind wing, tuberculated ridges of the |
fishbone pattern of the hind femur, and the appearance of the apex of phallus. As |
to the latter, especially in kevani, resemblance with the apex of phallus in Cranae |
patagiata Stal (type-species of that genus) is obvious.

Cranaella willemsei Ramme, 1941 |
(figs. 6—9, 14—15, 22, 28, pl. 1 fig. 4) |

Cranaella willemsei Ramme, 1941: 95, 218; C. Willemse, 1956: 108, 109.

Material studied: Surigao, Philipp. 14 (NMM), 19 (ANSP) (topotypes); P.I.,
Dinagat, Panamana, 20.iii.1945, H. H. Roberts 1 (ANSP).

Redescription. |

&, pl. 1 fig. 4. Differs from tuberculata as follows. Integument finely pitted,
impressions weaker and smaller. Head slightly longer. Prosternal process thick,
strong, roughly cubical, slightly curved anteriorly, basally slightly narrower than

F. WILLEMSE: The genus Cranaella 117

24

Figs. 20—24. Cranaella species, &, supra-anal plate and hind margin of last abdominal tergite, dorsal
view: 20, tuberculata Ramme, Surigao; 21, kevani sp.n., holotype; 22, willemsei Ramme, Surigao; 23,
rammei sp.n., paratype; 24, samarensis sp.n., holotype.

apically, anterior side higher than posterior one, ventral side quadrate and
flattened obliquely. Elytron wider, about three times as long as wide, hind margin
almost straight, venation extending over a larger area. Hind wing about three
quarters length of elytron, with some indistinct veins. Ridges of the fishbone
pattern of the outer medial area of the hind femur prominent but without
tubercles. Outer carinulae of hind femur slightly tuberculate. Hind margin of last
abdominal tergite thickened, widely and weakly emarginate in the middle and with
no more than an indication of a pair of furculae (fig. 22). Supra-anal plate longer
than wide (fig. 22). Phallic complex as in figs. 6—9. Apex of phallus (figs. 14—15)
with the ventro-posterior side slightly flattened and the tip divided into a narrow,
short, medio-distal process and a stronger, auricular, lateral lobe, which is curved
dorso-anteriorly.

Coloration about as in tuberculata, differing mainly in the ridges of the outer
medial area of the hind femur which are black, sharply contrasting with the
brownish colour of the hind femur.

©. Larger than the male. Subgenital plate (fig. 28) slightly longer than wide,
ventrally slightly depressed in the middle, keels moderately developed, hind
margin slightly tridentate, the median point more extending posteriorly than the
obtuse lateral ones. Coloration as in the male.

Measurements (length in mm): body, & 25.0-26.1, Q 28.5-31.4; pronotum,
& 4.4-4.5, 9 5.0-5.8; elytron, & 4.0-4.2, © 6.4-6.7; hind femur, & 12.5-13.4,
Q 14.3-14.5.

Distribution: Philippine Is. (Mindanao; Dinagat).

118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 4, 1977

Discussion. The species is well-defined by the male genitalia. The position
among the other species of the genus will be discussed under samarensis.

Cranaella rammei sp.n
(figs. 16—17, 23, 29, pl. 1 figs. 5—6)

Material studied: ¢ holotype, 9 allotype, 2¢ 19 paratypes, labelled: Dagami,
Leyte, P.I., Mt. Lobi, 10(1¢) & 21 (18) & 25.vi (allotype) & 4 (19) & 10.viii.1945
(holotype), E. R. Helwig (ANSP).

Description.

&, pl. 1 fig. 5. Differing from willemsei as follows. General appearance more
robust. Hind margin of last abdominal tergite (fig. 23) with a narrow median
incision, bordered laterally by a pair of narrow, obtusely pointed furculae. Apex of
phallus more elongate, distal process longer and lateral lobe moderately developed
(figs. 16—17).

Coloration as in willemsei. Posterior veins of elytron sometimes yellowish white.

©, pl. 1 fig. 6. Larger than the male. Subgenital plate (fig. 29) more elongate
than in willemsei, hind margin more produced posteriorly. Coloration as in the
male.

Measurements (length in mm): body, & 24.2-25.1, 9 29.0-34.1; pronotum,
Sd 4.4-4.5, 9 5.8-6.0; elytron, & 4.5-5.2, 9 5.9-6.1; hind femur, & 13.3-14.2,
Q 16.7-17.0.

Distribution: Philippine Is. (Leyte).

Discussion. The species is well-defined by the male genitalia. There are no
previous records. Its position among other species will be discussed under
samarensis.

Cranaella samarensis sp.n.
(figs. 18—19, 24, pl. 1 fig. 7)

Material studied: ¢ holotype, labelled: Island Samar, Baker (ANSP).

Description.

3, pl. 1 fig. 7. Much as willemsei, differing in abdominal terminalia. Furculae of
hind margin of last abdominal tergite large, triangular, apex pointed (fig. 24). Apex
of phallus with distal process and lateral lobe intermediate between those in
willemsei and rammei (figs. 18—19). Coloration as in willemsei.

©. Unknown.

Measurements (length in mm): &, body 26.2, pronotum 4.4, elytron 4.9, hind
femur 13.1.

Distribution: Philippine Is. (Samar).

Discussion. The species is defined by the furculae and the apex of phallus.
Together with willemsei and rammei, the three species form a species-group charac-
terized by the shape of the prosternal process, hind wings, apex of phallus, and

F. WILLEMSE: The genus Cranaella 119

28

26

27 29 30

Figs. 25—30. Cranaella species, 9, subgenital plate, ventral view: 25, carnipes Ramme, holotype; 26,
tuberculata Ramme, Surigao; 27, kevani sp.n., allotype; 28, willemsei Ramme, topotype; 29, rammei
sp.n., paratype; 30, multicolor Kevan, holotype.

black fishbone pattern of the hind femur. Both the hind wing, as well as the less
pitted integument and the shape of the apex of phallus, are intermediate between
the tuberculata-kevani group of Cranaella and some species of Cranae.

Cranaella multicolor Kevan, 1966
(fig. 30)

Cranaella multicolor Kevan, 1966: 41, pl. 3 figs. c—d.

This species was described after a single female from La Lun Mts., Davao Prov.,
Mindanao.

Marked characters may be summarized here: Pronotum with the posterior
margin of the dorsum straight; posterior margin of the lateral lobe not sigmoid but
straight and the lower posterior angle not produced; first transverse sulcus lacking
on the dorsum and the fourth one placed more anteriorly. Prosternal process
vertical and widened laterally. Elytron wide, about two-and-one-half times as long

120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 4, 1977

as wide, posterior margin nearly straight, venation reaching the margins. Hind
wing vestigial. Fishbone pattern of outer side of hind femur not tuberculate. Hind
margin of subgenital plate strongly tridentate, the lateral points as long as and
wider than the medial point (fig. 30).

Coloration. Head yellow with chocolate-brown stripe over occiput and behind
eyes. Pronotum chocolate-brown. Elytron green, yellowish at base. Hind femur
ochreous yellow, suffused blue-black in basal third, carinulae and fishbone pattern
unicolorous. Hind knee without ante- or postgenicular rings.

Measurements (length in mm): ©, body 28.5, pronotum 5.0, elytron 6.5, hind
femur 19.5.

Distribution: Philippine Is. (Mindanao).

Discussion. The shape and venation of the elytron, the shape of the pronotum
and the coloration disagree with other members of Cranaella. Its placement in this
genus is doubtful. Pronotum and elytron agree rather with Paracranae C. Willemse
from Celebes. However, the hind margin of the female subgenital plate and the
fastigium verticis are different in that genus. A similar shape of elytron is found in
Cranae luctuosa C. Bolivar and Cranae kuekenthali Brunner, both from the
Moluccas. As the male of multicolor is unknown, a more precise allocation of the
species has to be postponed.

REFERENCES

Kevan, D. K. McE., 1966. Some Orthoptera - Caelifera from the Philippine, Bismarck and Solomon Is-
lands, with a few interesting records from New Guinea and the Moluccas. — Ent. Meddr. 34:
375—420, figs.

Ramme, W., 1941. Beitrage zur Kenntnis der Acrididen-Fauna des indomalayischen und benachbarter
Gebiete (Orth.). — Mitt. zool. Mus. Berl. 25: 1—243, figs.

Willemse, C., 1956. Synopsis of the Acridoidea of the Indo-Malayan and adjacent regions (Insecta,
Orthoptera). Part II. Fam. Acrididae subfam. Catantopinae. Part one. — Publties natuurh. Ge-
noot. Limburg 8 (1955): 1-226, figs.

A STUDY ON THE GENUS CRANAE STAL
(ORTHOPTERA, ACRIDOIDEA, CATANTOPINAE)

by

FER WILLEMSE
Laurastraat 67, Eygelshoven, The Netherlands
With 45 text-figures and five plates

ABSTRACT

In Cranae at least 20 species are recognized. Unlike in previous studies, the diagnostic emphasis is
laid on morphological distinction rather than on colour characters. Cranae rufofemorata Willemse, and
its variety obscura, are synonymized with Cranae tibialis Brunner. Cranae patagiata coerulipes (C.
Bolivar) is considered a synonym of Cranae patagiata Stal. The following new species are described:
genjam, manokwari, longipennis, rubra and glabra. Some species, preliminarily allocated to Cranae, link
the genus along various lines with the genera Cranaella and Opiptacris.

INTRODUCTION

Cranae and some related genera form a large, natural group of species, each
with a very restricted range. Few characters have been used for the generic
distinction. When more characters are analyzed simultaneously, the present
generic classification appears largely artificial and certainly does not reflect a
hypothetical phylogeny. Several species, along various lines, link the discon-
tinuities between the genera. A generic revision is needed but material is scarce
and much basic information on the species concerned is wanting. Therefore, it
appears sensible to retain Cranae preliminarily as a taxonomic unit in its present
state. Depositories of the material used in this study are given in abbreviated form
throughout the text:

ANSP — Academy of Natural Sciences of Philadelphia
BPBM — Bernice P. Bishop Museum, Honolulu

ITZ — Instituut voor Taxonomische Zoölogie, Amsterdam
MC — Macdonald College, Quebec, Canada

MNHN — Museum National d'Histoire Naturelle, Paris
NMM — Natuurhistorisch Museum, Maastricht

NMW W— Naturhistorisches Museum, Wien

RNH — Rijksmuseum van Natuurlijke Historie, Leiden

ZMHU — Zoologisches Museum der Humboldt-Universität, Berlin

121

122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

Plate 1. Figs. 1—7. Cranae species, face, frontal view, similar scale: 1, patagiata Stal, 9 (type of C.

pervittata Brunner); 2, unistrigata (De Haan) Form 1, 9 (Aindoea R.); 3, nigroreticulata Brunner, &

(Tuguaer-Tasso); 4, longipennis sp.n., 9 (allotype); 5, luctuosa C. Bolivar, 9 (Telega, Obi I.); 6, glabra

sp.n., Q (paratype); 7, pictipennis C. Willemse, 9 (paratype). Fig. 8. C. patagiata Stal, & (type of C.
pervittata Brunner), pronotum, dorsal view.

F. WILLEMSE: The genus Cranae 123

My thanks are due to the following persons: J. L. Gressitt and the late Miss S.
Nakata, Honolulu; M. Beier, Vienna; K. K. Günther, Berlin; P. H. van Doesburg,
Leiden; M. Descamps, Paris; D. K. McE. Kevan, Macdonald College;
H. Radclyffe Roberts and D. Rentz, Philadelphia; and especially to C. Jeekel,
Amsterdam, for reading the manuscript and for his critical comments.

Cranae Stal, 1878

Cranae Stal, 1878: 41, 85; Brunner, 1893: 135; 1898: 236; Kirby, 1910: 387; C. Willemse, 1921: 7, 21; C.
Bolivar, 1932: 393; Uvarov, 1937: 17; C. Willemse, 1939: 74; Ramme, 1941: 87; C. Willemse, 1956:
9,97.

Type-species: Cranae patagiata Stal, 1878.

A diagnosis, based on the revised material, reads as follows:

Medium size. Integument shiny. Face and pronotum more or less pitted or
almost smooth, face occasionally more or less wrinkled. Occiput, on either side,
with a row of slight transverse impressions.

Antennae filiform, segments up to five times as long as wide, tip reaching nearly
middle of hind femur or slightly longer. Head thick, round. Eyes ovoid-
hemispherical. Interocular distance as wide as or a little narrower than greatest
width of fastigium verticis. Distance between eye and lower margin of cheek much
shorter than vertical diameter of eye. Fastigium verticis at lower level than vertex
between eyes, usually slightly marked off from rest of vertex; sometimes with fine
median sulcus and fissured apex, merging with sulcus of frontal ridge; more or less
widely triangular from above, truncated apex more or less wide and not reaching
tip of scape; in profile, from subhorizontal to distinctly declivous, about
rectangularly meeting face. Face (pl. 1 figs. 1—7) slightly reclinate. Frontal ridge
present only above the median ocellus, weakly sulcate; margins obtuse, nearly
parallel, sometimes connected transversely at median ocellus. Lateral facial keels
straight, low, more or less divergent; face, along these keels, depressed in some
species. Occiput and cheeks convex.

Pronotum (pl. 1 fig. 8, pl. 2 fig. 9—16) about as long and as wide as head or
slightly shorter or narrower, without keels, cylindrical or dorsum slightly
depressed from above, lateral lobes parallel or slightly compressed laterally in the
middle. Four transverse sulci, more or less deep, second one dorsally only.
Dorsum divided by the sulci into five parts; width of these parts varying
individually, but distance between second and third sulci usually slightly smaller,
that between third and fourth sulci usually slightly larger. Lateral lobe about as
long as high or shorter; deepest point at level of third sulcus, from there lower
margin concave anteriorly, straight or slightly convex posteriorly; anterior angle
obtuse-angulate, posterior angle about rectangular, from narrowly rounded to
obtusely pointed. Anterior margin of dorsum weakly rounded, of lateral lobe
straight and divergent anteriorly. Posterior margin of dorsum weakly rounded,
straight or weakly emarginate, of lateral lobe straight or slightly concave, vertical
or slightly slanting upwards posteriorly. Prosternal process strong, more or less

124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

Plate 2. Figs. 9—16. Cranae species, pronotum, dorsal view, similar scale: 9, tibialis Brunner, &

(holotype); 10, nigroreticulata Brunner, & (Tuguaer-Tasao); 11, unistrigata (De Haan), Form |, ¢ (Fak

Fak, T.C. Maa); 12, trivittata C. Willemse, © (Bivak Eiland); 13, rufipes Ramme, © (Finschhafen); 14,

luctuosa C. Bolivar, & (Telaga, Obi I.); 15, glabra sp.n., 9 (paratype); 16, pictipennis C. Willemse, &
(allotype).

F. WILLEMSE: The genus Cranae 125

laterally compressed, in profile vertical or slightly directed anteriorly; truncated
apex widened laterally, its anterior margin straight and posterior margin curved
towards anterior one. Mesosternal lobes wider than long, inner margins convex,
interspace wider than long, about as wide as a lobe. Metasternal interspace
narrowly triangular. Mesonotum about two-thirds of metanotal length. Meta-
notum narrower than distance between anterior margin of pronotum and second
pronotal sulcus.

Plate 3. Figs. 17—24. Cranae species, spread left elytron, outer view, similar scale: 17, patagiata Stal, 9

(‘Java’); 18, unistrigata (De Haan), Form 1, 9 (Aindoea R.); 19, manokwari sp.n., 9 (allotype); 20,

longipennis sp.n., 9 (allotype); 21, kuekenthali Brunner, 9 (Halmaheira, T. Barbour); 22, luctuosa C.

Bolivar, 9 (Telaga, Obi I.); 23, glabra sp.n. 9 (paratype); 24, pictipennis C. Willemse, @ (holotype) (in
situ)

126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

Tegmina (pl. 3 fig. 17—24) from squamipterous to subbrachypterous, ranging
from reaching first up to ninth abdominal tergite. Elytra touching each other or
more or less separated dorsally; from almost as long as wide to three times as long
as wide; posterior margin from straight to strongly rounded; anterior margin more
or less evenly rounded or tapering in apical half; apex, at the end of folding of
elytron, divided into a larger anterior and a smaller posterior lobe (except glabra
and pictipennis); venation simple, a wider zone along the anterior and a much
narrower one along posterior margin without veins. Hind wing slightly shorter
than elytron, with sparse veins (as a small, membraneous fold in pictipennis).
Tympanum open, almost circular, level with the body surface (as a closed, narrow
furrow in pictipennis).

Legs stout (attenuate in glabra). Hind femur about reaching tip of abdomen
(male), or slightly shorter (female); fishbone pattern distinct and smooth, keels
smooth or slightly crenulate, inner lower carinula pilose. Knee-lobes truncate.
Brunner’s organ present. Hind tibia slightly shorter than hind femur; margins
rounded, strongly pilose; six or seven inner and four or five outer spines regularly
arranged, apical ones included. Hind tarsus pilose, slightly longer than half the
tibial length, third segment longest, first and second segments of about equal
length.

Male. — Hind margin of last abdominal tergite ( fig. 5) triangularly excised with
a pair of well-separated, small, widely triangular furculae. Supra-anal plate (fig. 5)
wider than long, roughly triangular; lateral margins thickened near base and there
with a short, low, transverse ridge, extending medially; middle of basal half
depressed; apex more or less widely parabolic. Cercus slender, slightly up- and
incurved, pointed apex reaching or extending beyond tip of subgenital plate. The
latter subconical, short and its apex obtuse.

Phallic complex, figs. 6—45. Epiphallus narrowly bridge-shaped; ancorae, in
most species, more (fig. 7) or less (fig. 14) joined with lateral plates, in few
species present as a separate projection (fig. 11); inner lophi smaller and tooth-
shaped, outer lophi larger and hook-shaped. Ectophallic membrane with shield-
shaped dorsal sclerite. Cingulum with narrowly U-shaped apodemal structure.
Rami extending latero-ventrally and dorso-posteriorly; usually narrow (fig. 9), but
in some species wider (fig. 13). Cingular valves short, not by far reaching tip of
apical penis valves, triangular from above. Apical penis valves slightly upcurved
and joining laterally with sheath of penis. Apex of phallus formed by sheath of
penis and projecting ends of apical penis valves. Shape of apex of phallus variable
but species-specific (figs. 16—45).

Female. — Hind margin of last abdominal tergite triangularly excised. Supra-
anal plate tongue-like, twice as long as wide, apex parabolic. Cercus conical,
slightly outcurved, not reaching tip of supra-anal plate. Subgenital plate (fig. 1)
twice as long as wide, ventral surface flattened in the middle and, usually, rather
depressed or sulcated apically between a pair of short, obtuse, longitudinal keels;
hind margin slightly sigmoid laterally, tridentate in the middle, median point
longest and widest, lateral points corresponding with distal ends of ventral keels.
Dorsal aspect of subgenital plate (fig. 3) with a pair of simple, round columellae.

F. WILLEMSE: The genus Cranae 127

Valves of ovipositor (fig. 2) slender, straight, margins finely serrate or almost
smooth, apices not definitely hooked. Lateral basivalvular sclerite elongate-
triangular, ventral basivalvular sclerite three to four times as long as wide.
Spermatheca (fig. 4) simple, with an apical diverticulum and a wider, strongly
curved, preapical diverticulum.

Coloration. Most species characteristically coloured. Sexual dichromatism not
apparent. Individual variation appearing to be slight, but a general statement not
possible by lack of material. Coloration of the following parts similar in all taxa of
the genus: eyes, individually, from yellow to dark brown; apical part of mandible
black; crescents of hind knee dark castaneous brown; spines of hind tibia and
claws of tarsus dark brown or blackish, tips black; furculae in the male black.

General distribution: New Guinea and the Moluccas.

Discussion. Previously, ten species and three subspecies have been placed in
Cranae (C. Willemse, 1956). Now at least 20 species are recognized. Opiptacris
pictipennis (C. Willemse) has been re-established to its original combination Cranae
pictipennis (F. Willemse, 1975: 121). Cranae rufofemorata C. Willemse and its
variety obscura are both considered conspecific with Cranae tibialis Brunner.
Cranae patagiata coerulipes (C. Bolivar) is considered conspecific with Cranae
patagiata Stal. The available material clearly shows that Cranae unistrigata (De
Haan) and trivittata C. Willemse belong to a complex of closely related
populations. Two of these have been described by Ramme, as caprai and trivittata
rufipes, respectively. Whether caprai should be considered taxonomically distinct
from nominate unistrigata is an open question. However, rufipes is now given
specific rank. As to the remaining material resembling nominate unistrigata and
trivittata, it is still hard to decide whether speciation has been completed or not. In
two cases only differences seem sufficient for distinction on species level: genjam
and manokwari. Eight additional species are recognized as new. Three are fully
described and named: /ongipennis, rubra and glabra. The others could not be
defined precisely enough, by lack of males. They are briefly recorded, but not yet
named.

In Cranae, some natural species groups can be distinguished. The main stock of
the genus is found on New Guinea. It is formed by the unistrigata complex. The
following groups among Cranae may be recognized: (1) nominate unistrigata,
genjam, and (probably) the not fully known species from Roon I. and Sabang; (2)
trivittata, rufipes, manokwari and (probably) the not fully known species from Bubia
and Normanby I.; (3) rubra; (4) longipennis and (probably) the not fully known
species from Aseki; (5) patagiata, tibialis; (6) nigroreticulata; (7) species with
important discontinuity as kuekenthali, luctuosa, glabra, and pictipennis. Especially
the latter group of species links, through various characters, Cranae with Cranaella
Ramme and Opiptacris Walker.

Key to the species of Cranae (emendata Brunner and the now recognized but not
yet named new species omitted):

1. Elytron circular and not reaching beyond the first abdominal tergite (pl. 3 fig.
24); tympanum closed (Buru) ......... pictipennis C. Willemse (p. 149)

128

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

Elytron elongate and reaching beyond the first abdominal tergite (pl. 3 figs.
1723) !tympanumlopen), circular. in. RR EEE À

. Elytron unicolorous olivaceous green, without black (Keils.) ..........

ML 2 DISS REDE ARR AM EN STAR SP ENO A tibialis Brunner (p. 132)
Elytron with at least the anterior margin and the apex bordered with black or

dark brown Wera DE LH L'ATRIO DINI TENEN 3)

» Blytron'red'and'black:!. „00 Mt IRE SARO A I 4
Elytron without red … ‚no orn Je ORNE EERE 5

. Hind femur red on basal half (Woodlark I) ........ rubra sp.n.(p. 144)
Hind femur not red (Ambon; Ceram) .......... patagiata Stal(p. 128)

. Elytron not reaching beyond the second abdominal tergite (pl. 3 fig. 23)
(Biak LE a er LEE EE RIO ER glabra sp.n. (p. 147)
Elytron longer (pl. 3fig: 1722). wa: WAE ENE 6

. Pronotum cylindrical, without longitudinal fasciae (pl. 2 fig. 14); phallic
complex with the rami wide (fig M2) =) ae = 7
Pronotum slightly depressed from above, with longitudinal fasciae (pl. 2 figs.
10—13); phallic complex with the rami narrow (fig. 9) ............ 8

. Hind femur orange on basal half (Halmahera) .. kuekenthali Brunner (p. 145)
Hind femur not orange on basal half (Obi I.) .. /uctuosa C. Bolivar (p. 146)

. Median black fascia of pronotal dorsum rhomboidally widened in the middle
(piste 10) (Halmaheira) Demenz nigroreticulata Brunner (p. 135)
This fascia not widened in the middle (pl. 2 figs. 11—13) ........... 9

. Elytron wide and long, margins about parallel, the yellow streak wide (pl. 3 fig.
20) (Ss New Guinea) Ks EI 0 longipennis sp.n. (p. 142)
Elytron shorter, tapering apically, the yellow streak narrow (pl. 3 figs. 18—19)
i ER Se mere aera RN NIUE RR AA o 5 0 0 0 0 10

. Yellow antegenicular ring of hind femur bordered proximally with red or
orange. Sal e kN LR RE Ue RO Rr 11
This ring bordered proximally with a narrow black ring, either complete or not
e a ll u eM Eelen 0 © © oo oo 0 12

. Apex of phallus short (figs. 22—23) (Genjam, near Hollandia) ..........

Re ENS ne ICI ME (SU INTER BETEN genjam sp.n. (p. 140)
Apex of phallus elongate (figs. 24—31) (W & N. New Guinea) ..........
Latte. RE ro NE MEURT RE AS unistrigata De Haan (p. 135)

. Pronotum shorter, dorsum more depressed (pl. 2 fig. 12); head more globose

(SW:NewiGuinea) ra AR INS. trivittata C. Willemse (p. 140)
Pronotum longer, dorsum less depressed (pl. 2 fig. 13); head less globose (N.
New Guinea) mite rufipes Ramme (p. 141) and manokwari sp.n. (p. 142)

Cranae patagiata Stal, 1878
(figs. 1—2, 5—10, 16—17, pl. 1 figs. 1, 8, pl. 3 fig. 17, pl. 4 fig. 25)

Cranae patagiata Stal, 1878: 85; Brunner, 1898: 236; Kirby, 1910: 387; Sjöstedt, 1932: 62; C. Willemse,

1939: 74: Ramme, 1941: 87, 88; C. Willemse, 1956: 104 (not Brit. N. Guinea).

Cranae patagiata patagiata; C. Willemse, 1956: 99.

F. WILLEMSE: The genus Cranae 129

Plate 4. Figs. 25—30. Cranae species, lateral view: 25, patagiata Stal, 3 (type of ©. pervittata Brunner);

26, tibialis Brunner, & (paratype of C. rufofemorata ©. Willemse); 27, unisirigaia (De Haan), &

(holotype); 28, genjam sp.n., & (holotype); 29, manokwari sp.n., ¢ (holotype); 30. luctuosa C. Bolivar, 3
(Telaga, Obi L.).

130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

Cranae pervittata Brunner, 1898: 236, 238; Kirby, 1910: 387; C. Willemse, 1922: 716; C. Bolivar, 1923:
146: C. Willemse, 1939: 74; Ramme, 1941: 88 (synonymy established); C. Willemse, 1956: 104 (in sy-
nonymy only).

Cranae pervittata pervittata; C. Bolivar, 1923: 147; C. Willemse, 1939: 74.

Cranae pervittata coerulipes C. Bolivar, 1923: 147; C. Willemse, 1939: 74. Syn. nov.

Cranae patagiata coerulipes: Ramme, 1941: 88 (nec coerulescens); C. Willemse, 1956: 99, 105; Kevan,
1966: 411.

i
N
|

Material studied: & lectotype, © paralectotype of Cranae patagiata, labelled: Col. |
Br. v. W. Amboina, 1258, det. Br. v. W. Cranae patagiata (NMW). Both specimens |

discoloured.
Types of Cranae pervittata, 13 19, labelled: Coll. Br. v. W. Amboina

Staudinger, 14.141, Det. Br. v. W. Cranae pervittata, Type, Cranae patagiata Stal

Ramme det. 1939 (NMW).

Additional material: Amboina, Dr. Doleschal, 1859 (2g, 29, discoloured)
(NMW); Molukken, Depuiset ded., 5043, det. Br. v. W., Cranae pervittata COR
(NMW); Amboine, Moluques, Collection Brunner, Cranae patagiata Stal, collec-
tion A. Finot (14, 19, discoloured) (MNHN); Amboine, Collection A. Finot,
Cranae pervittata Br. (14) (MNHN); Ceram, Piroe (13) (ZMHU); Ceram, Piru,
F. Barbour, 1906-07 coll. (19, discoloured) (ANSP); Java, Oberth., ex coll.
Bolivar, Cranae patagiata, det. C. Willemse (1 9) (NMM).

Redescription.

Male, pl. 4 fig. 25. Face and thorax slightly pitted. Interocular distance less than
greatest width of fastigium verticis. Fastigium verticis triangular, greatest width
and length about equal, finely sulcate apically, in profile slightly declivous. Frontal
ridge with low margins, below median ocellus disappearing into slightly wrinkled
surface of face. Face slightly transversely convex, weakly impressed along lateral
keels.

Pronotum (pl. | fig. 8) as long and as wide as head, slightly depressed from
above, lateral lobe parallel; sulci moderate; posterior margin of dorsum slightly
extending posteriorly, straight or very slightly rounded; lateral lobe as long as high,
posterior angle narrowly rounded, posterior margin straight and vertical.

Elytron reaching middle of hind femur, about three times as long as wide,
posterior margin straight, anterior margin strongly rounded basally, tapering
apically towards the widely rounded, divided apex. Both elytra touching each
other apically, narrowly separated basally.

Supra-anal plate (fig. 5) with the apex widely parabolic. Furculae (fig. 5) very
small, well separated. Cerci reaching tip of subgenital plate. Phallic complex, figs.
6—10, 16—17. Ancorae of epiphallus joined with lateral plates. Cingular rami
narrow. Apex of phallus with moderately developed sheath of penis and short,
strongly tapering apical penis valves, their tips almost pointed.

Coloration olivaceous yellow, dark bluish-black and red. Antennae dark brown,
paler apically. Head and pronotum olivaceous yellow with three bluish-black
fasciae: on either side, from behind eye over middle part of pronotal lateral lobe
and a median fascia from between the eyes over occiput and pronotal dorsum;
margins of fastigium verticis and frontal ridge, lateral facial keels and larger or

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F. WILLEMSE: The genus Cranae 131

Figs. 1—2. Cranae patagiata Stal, © (type of C. pervittata Brunner): 1, tip of abdomen, ventral view; 2,

same, lateral view. Figs. 3—4. Cranae unistrigata (De Haan), Form 6, 9 (Hollandia): 3, subgenital plate,

dorsal view; 4, spermatheca. Figs. 5—10. Cranae patagiata Stal, & (type of C. pervittata Brunner): 5,

furculae and supra-anal plate; 6, epiphallus, dorsal view; 7, same, posterior view; 8, phallic complex.

dorsal view, epiphallus and part of ectophallic membrane removed; 9, same, lateral view; 10, same,
endophallus.

132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

smaller areas of face, bluish or greenish-black; episternum, meso- and metathorax
blackish, except for yellowish lower part of second episternum and epimerum.
Elytron dull red, anterior margin and apex widely bordered with black. Hind
wing infumate. Abdomen violaceous brown and yellowish. Fore and middle
femora dull red, apical margins of knees, whole tibiae and tarsi olivaceous green
and yellow. Hind femur ranging from violaceous black to yellowish; in the former
case, dorsal side yellowish, in the latter one, carinulae and fishbone pattern black.

Antegenicular part of hind femur, as long as length of hind knee, yellow. Hind _

knee, tibia and tarsus dark bluish-green; hind tibia with a narrow, indistinct,
greenish postgenicular ring; hind tarsus paler below.

Female. Larger than male. Face, pl. 1 fig. 1. Elytra (pl. 3 fig. 17) more widely
separated dorsally and reaching distal end of basal third of hind femur, about two-
and-one-half times as long as wide. Abdominal terminalia, figs. 1—2. Subgenital
plate distinctly sulcate apically. Coloration as in male.

Measurements (length in mm): body, & 22.0-23.0 9 27.0—30.0; pronotum, ¢
4.2—4.5, 9 5.2—5.8; elytron, & 8.2—8.8, 9 7.6—8.9; hind femur, ¢ 11.7—12.2, 9
13.6— 14.9.

Distribution: Moluccas (Ambon, Ceram).

Discussion. The species is well-defined, especially by the phallic complex and
the coloration. The apex of phallus comes close to that of tibialis and genjam. The
coloration of the elytron is much as in rubra.

The discoloured pair, bearing Brunner’s identification label patagiata, lacks any
type indication. In agreement with the original paper, the record by Sjöstedt (1932)
and Ramme’s opinion (1941), the pair should be considered Stal’s types. The male
is hereby designated lectotype. Ramme synonymized Brunner’s pervittata with
Stal’s patagiata. Comparison of the phallic complex definitely reveals their
synonymy. Bolivar’s pervittata coerulipes from Ceram is defined by slightly distinct
coloration. However, this character is not reliable as among patagiata material
from Ambon (MNHN) a transitional form is also found. Besides, genitalia and
other characters are very similar and the localities quite close. Bolivar’s coerulipes
is considered conspecific with patagiata.

The previous records from British New Guinea (material not at hand) and Java
(1 © at hand) are probably incorrect. As to the former, see below under rubra.

Cranae tibialis Brunner, 1898
(figs. 18—19, pl. 2 fig. 9, pl. 4 fig. 26)

Cranae tibialis Brunner, 1898: 236, 237; Kirby, 1910: 287; C. Willemse, 1939: 74; 1956: 98, 99.
Cranae rufofemorata C. Willemse, 1939: 73, 74, fig. 1; 1956: 98, 100, fig.: F. Willemse, 1966a: 38; 1966b:

65. Syn. nov.

Cranae rufofemorata var. obscura C. Willemse, 1956: 98, 101; F. Willemse, 1966a: 38; 1966b: 65. Syn.
nov.

Material studied: ¢ holotype of Cranae tibialis, labelled: Coll. Br. v. W. Key-
Inseln Ribbe leg., det. Br. v. W. Cranae tibialis, Type (NMW). The specimen lacks
both antennae; right tegmina are spread.

Type-series of Cranae rufofemorata, labelled: H. C. Siebers Kei Eil., Gn. Daab

F. WILLEMSE: The genus Cranae 133

Figs. 11—13. Cranae kuekenthali Brunner, & (paratype): 11, epiphallus, dorsal view; 12, phallic

complex, dorsal view, epiphallus and part of ectophallic membrane removed; 13, same, lateral view.

Figs. 14—15. Cranae longipennis sp.n., & (holotype): 14, epiphallus, dorsal view; 15, same, posterior

view. Figs. 16—21. Cranae species, 3, tips of apical penis valves and sheath of penis, ventro—posterior

(even numbers) and left lateral (odd numbers) view, similar scales: 16—17, patagiata Stal (type of C.

pervittata Brunner); 18—19, tibialis Brunner (paratype of C. rufofemorata C. Willemse); 20—21,
nigroreticulata Brunner (Tuguaer—Tasao).

134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

143 (¢ holotype) & 138 (14 paratype) & Toeal 201 (14 paratype, 9 allotype),
Cranae rufofemorata n.sp. C. Willemse det. (Holo- and allotype RNH, paratype |
NMM). The holotype lacks both antennae.

Type-series of Cranae rufofemorata var. obscura, labelled: H. C. Siebers Kei Eil |
Gn. Daab 85 (9 holotype) & 135 (9 paratype), Cranae rufofemorata Will. var.
obscura n.var. Det. C. Willemse (NMM). The holotype lacks the right antenna and |
both fore legs.

Redescription

Male, pl. 4 fig. 26. Differs from type-species in pronotum, apex of phallus and |
coloration. Pronotum (pl. 2 fig. 9) more cylindrical, slightly wider, sulci slightly |
deeper, posterior margin extending less posteriorly and lateral lobe narrower. |
Apex of phallus (figs. 18—19) slightly larger, sheath of penis more developed, |
apical penis valves more narrowing apically and tips larger.

Coloration as described by C. Willemse (1956) under rufofemorata and its variety | |
obscura. Marked characters may be summarized as follows: head from yellowish or |
yellowish-green to almost completely dark olivaceous; pronotum from yellowish- |

elytron completely yellowish or greenish; legs red, except for hazel brown hind |
knees with blackish lobes, a yellowish-green antegenicular ring of hind femur, |
dark brownish or violaceous apical third of hind tibia, and brownish or greenish | |
tarsi.

Female. Larger than male. Abdominal terminalia as in type-species. Coloration |
as in male, except for hind femur, which is yellowish-green with lower inner |
marginal area red. |

Measurements (length in mm): body, & 21.0—22.0, © 31.0— 35.0; pronotum, & | |
4.0—4.3, 9 5.2—5.4; elytron, & 7.7—8.0, 9 7.5—8.2; hind femur, Z 11.3—12.5, 2
14.2—15.2. |

Distribution: Moluccas (Kei Is.). |

Discussion: The species is well-defined, especially by the apex of phallus and the
coloration. Apex of phallus much as in patagiata and genjam. The unicolorous
elytra without black forming a particular feature of the species. Shape and
coloration of the pronotum intermediate between patagiata and the kuekenthali-
luctuosa group.

As against the original statement and in agreement with the given measure-
ments, the holotype is not a female but a male. Comparison of the holotype of
tibialis with the males of the type-series of rufofemorata, clearly reveals their
synonymy. C. Willemse distinguished among rufofemorata, the variety obscura. It
was based on the dark olivaceous green head and pronotum in two females of the
type-series of rufofemorata. However, this character is not reliable, as the colour of
head and pronotum in males of nominate rufofemorata ranges from pale olivaceous
green to as dark as in obscura. The darkest male of nominate rufofemorata is the
holotype (RNH). C. Willemse’s distinction may be explained by assuming that this
holotype was not before him when he described the variety (1956). By lack of
distinct features, I consider obscura conspecific with rufofemorata as well as with
tibialis.

F. WILLEMSE: The genus Cranae 135

Cranae nigroreticulata Brunner, 1898
(figs. 20—21, pl. 1 fig. 3, pl. 2 fig. 10)

Cranae nigroreticulata Brunner, 1898: 198, 236, 239; Kirby, 1910: 387; C. Willemse, 1922: 715; 1939: 74;
Ramme, 1941: 90; C. Willemse, 1956: 98, 101.

Cranae marginata Brunner, 1898: 198, 236, 237 (partim); Kirby, 1910: 387 (do); Ramme, 1941: 87, 90 (9
synonymy established); C. Willemse, 1956: 101 (in synonymy only).

Material studied: 1 3, labelled: Halmahera, Tuguaer-Tasoa, 100-150 m, 20 ix.
1951 (13) (NMM).

Redescription

Male. Differs from the type-species in head, pronotum, abdominal terminalia,
and coloration. Fastigium verticis narrowly triangular, apex not fissured, in profile
subhorizontal. Face (pl. 1 fig. 3) more pitted, not wrinkled, more transversely
convex, lateral keels lower. Frontal ridge weaker. Pronotum (pl. 2 fig. 10)
narrower, sulci slightly wider and posterior margin of lateral lobe slightly concave.

Furculae larger, more pointed and closer together. Supra-anal plate shorter,
apex wider and transverse ridges more distinct. Apex of phallus larger and more
elongate (figs. 20—21); sheath of penis narrow; apical penis valves wider and less
tapering apically.

Coloration as described by C. Willemse (1956). Marked characters may be
summarized as follows: head yellow; pronotum orange-yellow, dorsum with a
black median fascia, in the middle rhomboidally widened, lateral lobe with a black
fascia in upper part; elytron yellowish, anterior margin. narrowly, apex widely
bordered with black; fore and middle legs orange-yellow with some black
markings; hind femur orange-yellow with carinae, carinulae and fishbone pattern
black, antegenicular area completely orange-yellow; hind knee black; hind tibia
and tarsus violaceous black.

Female. Not studied.

Measurements were given in previous papers.

Distribution: Moluccas (Halmahera).

Discussion. The species is well-defined by the face, apex of phallus and
coloration. The pitted and not wrinkled face and the rhomboidal figure of the
pronotal dorsum are unique in Cranae. The apex of phallus agrees with unistrigata
rather than with patagiata.

Type-material of Cranae marginata was not at hand. No comments are given as
to the synonymic notes in Ramme (1941).

Cranae unistrigata (De Haan, 1842)
(figs. 3—4, 24—31, pl. 1 fig. 2, pl. 2 fig. 11, pl. 3 fig. 18, pl. 4 fig. 27)

Acridium (Oxya) unistrigatum De Haan, 1842: 143, 156, 158, 246, pl. 21 figs. 7—7a (g), 8 Guv. &, not 9);
I. Bolivar, 1918: 42 (juv. &, not 9); C. Willemse, 1928: 11.

Acridium unistrigatum; Stal, 1873: 53.

Oxya unistrigata; Walker, 1870: 648.

Oxya (?) unistrigata; Kirby, 1910: 395 Guv. &, not 9).

136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

Taeniophora unistrigata; Bruner, 1907: 245, 247. |
Chitaura haani I. Bolivar, 1918: 34 Guv. &, not 9); C. Willemse, 1956: 106 (in synonymy only). |
Cranae unistrigata; I. Bolivar, 1898: 88; Kirby, 1910: 387; C. Willemse, 1922: 716; 1932: 46; 1939: 75; |

Ramme, 1941: 87, 90; C. Willemse, 1956: 99, 106 (not Sabang); Kevan, 1966: 411 (not Roon I).

Material studied: & holotype, labelled: N. Guinea, Type, Cranae unistrigata | |
De Haan & Det. C. Willemse (RNH). The specimen lacks the left antenna and]
right fore leg.

Redescription of the holotype.
Male, pl. 4 fig. 27. Differs from the type-species in head, pronotum, elytron, |
abdominal terminalia and coloration. |
Head slightly more globose. Face wider. Lateral facial keels and margins of |
frontal ridge more raised. Interocular distance wider. Fastigium verticis more |
widely triangular, more declivous, and apex wider and not fissured. |
Pronotum (pl. 2 fig. 11) slightly wider, dorsum more depressed and sulci less |

Figs. 22—33. Cranae species, &, tips of apical penis valves and sheath of penis, ventro-posterior (even |
numbers) and left lateral (odd numbers) view, similar scales; 22—23, genjam sp.n. (holotype); 24—25,
unistrigata De Haan (holotype); 26—27, unistrigata (De Haan), Form 2 (Sorong); 28—29, unistrigata |
(De Haan), Form 4 (Jobi I.); 30—31, unistrigata (De Haan), Form 6 (Adelbert Mts.); 32—33, rufipes |

Ramme (?) (Toricelli Mts.). |

F. WILLEMSE: The genus Cranae 137

wide. Elytron narrower and longer, reaching just beyond middle of hind femur.
Furculae larger. Apex of phallus (figs. 24—25) longer and much more elongate;
apical penis valves narrow, very slightly tapering apically, tips obtuse.

Coloration olivaceous green, yellow and black. Antennae dark brown. Head
black except for olivaceous face and mouthparts, yellow fastigium verticis, a
yellow sharply delimited, ovoid spot on the occiput near the dorso-posterior
margin of each eye and a yellow narrow fascia from the antennal base along
anterior margin of eye and lower margin of cheek.

Pronotum yellow with three black fasciae: a median one over the dorsum, twice
as wide as each yellow lateral part of the dorsum and, on either side, over the
upper two-thirds of lateral lobe. Meso- and metathorax yellowish, except for first
episternum, upper part of pleurae and lateral parts of meso- and metasterna,
which are black. Elytron brownish-black with a narrow yellow streak along folding
of elytron. Hind wing slightly infumate. Abdomen brown, cerci and supra-anal
plate darker brown.

Fore and middle legs olivaceous green. Hind femur in basal half olivaceous
green, with fishbone pattern darker green, in distal half red with a yellow, narrow,
antegenicular ring; lower inner marginal area completely red; inner medial area
almost completely red. Hind knee, tibia and tarsus dark blue.

Measurements (length in mm): body, 19.0; pronotum, 3.8; elytron, 9.0; hind
femur, 11.5.

Distribution: New Guinea (SW coast).

Discussion. Contrary to Ramme’s (1941) statement, the holotype is in the
collection of the Leiden Museum (C. Willemse, 1928, 1956). The agreement of the
specimen with figure 7 in the original paper is conspicuous. The locality label
reads “N. Guinea”. In the original paper (p. 143) the locality is defined “Nieuw
Guinea’s zuid-westkust’’. The juvenile male of figure 8 in De Haan’s paper could
not be traced.

Comparison of the material at hand with the holotype of unistrigata reveals that
a complex of closely related forms has been referred to that species previously.
The differences in morphology and coloration are not conspicuous, but rather
stable. It must be pointed out that when the material from different localities
agrees in certain features, these localities always form a united section of the range
of the whole complex. Due to the scanty morphological differences, especially in
the shape of the apex of phallus, it is impossible to ascertain whether reproductive
isolation has been achieved by the various populations. Only in one case, the quite
distinct apex of phallus justifies the creation of a distinct taxon (genjam). The
situation in the unistrigata complex can be compared with that in Pseudocranae
bimaculata C. Willemse (F. Willemse, 1972: 53). I here arrange the material into six
distinct Forms, of which Form 1 agrees with the holotype and Ramme’s
“unistrigata Farbungsrasse Mt. Arfak’’, Form 2 with Ramme’s caprai, and Form 6
with Ramme’s “unistrigata Farbungsrasse Sepik-Gebiet”. The differences lie in the
projecting ends of the apical penis valves and in the coloration. As to the latter,
the yellow occipital spots, the width of the pronotal fasciae and the coloration of
the legs, especially of the hind femur, appear to be of taxonomic value.

138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

Form 1
Cranae unistrigata; Ramme, 1941: 90 (Färbungsrasse Mt. Arfak); C. Willemse, 1956: 107 (partim).

Material studied: Vogelkop, Fak Fak, S. Coast of Bomberai, 10—100 m, 12. vi.
1959, T. C. Maa (1 g) (MC); Fak Fak, C. J. L. Palmers (1 9, discoloured) (NMM);
S. W. New Guinea, S. coast, Kambrau near Argoniibaai, 4. viii. 1941 (19) & near
mouth of R. Aindoea, 12.viii. 1941 (19), E. Lundquist (NMM); Siwi (Forêt),
6.111.1929, Prince Léopold (1 ¢) (NMM); Siwi, Arfak, 21.iv.—25.v.1928, Mayr(1g,
1 9 ZMHU, 19 NMM).

The males of this material (all from Northwest New Guinea) agree fairly well
with the holotype of unistrigata, especially in the apex of phallus.

The female, not yet described for unistrigata (s.l.), is larger than the male. Face,
pl. 1 fig. 2. The tegmina are more separated dorsally and shorter than in the male.
The elytron (pl. 3 fig. 18) is two-and-one-half times as long as wide, anterior
margin moderately tapering apically. Abdominal terminalia as in the type-species.
Dorsal aspect of subgenital plate and the spermatheca as in figs. 3—4. Coloration
as in the male. Inner medial area of hind femur may be completely red, base of
lower inner marginal area dark olivaceous. The measurements agree with those
given by C. Willemse (1956). The Fak Fak material has not been recorded
previously.

Form 2

Cranae unistrigata; I. Bolivar, 1898: 88 (only Sorong); C. Willemse, 1939: 75 (do); Kevan, 1966: 411 (do).
Cranae caprai Ramme, 1941: 91, fig. 38; C. Willemse, 1956: 99, 107.

Material studied: Sorong, T. Barbour (1 &, 2 9) (ANSP); Nieuw Guinea
Expeditie 1903, Manikion, 14.—28.II (1 3) (NMM) (all discoloured).

Ramme described caprai after a single discoloured female from Sorong. The
present material from Sorong agrees with Ramme’s description. However, most, if
not all, distinct characters just may be due to shrivelling. The apex of phallus of
the Sorong male (figs. 26—27) differs from that of the holotype of unistrigata, in
shorter and more tapering apical penis valves with tips less obtuse. Whether this
distinction is reliable is difficult to evaluate as the specimen, like the female, is
shrivelled by spirit.

The present Sorong material has been referred to unistrigata by Kevan (1966),
the Manikion male to caprai by C. Willemse (1956). The locality of the latter could
not be traced. The apex of phallus of this male agrees with that of unistrigata's
holotype rather than with that of the Sorong male.

Form 3
Cranae unistrigata: 1. Bolivar, 1898: 88 (only Waigiou); C. Willemse, 1939: 75 (do); 1956: 106 (do).

Material studied: Waigeou, Staudinger (1 9) (NMM).
Material from Waigou Island has been referred to unistrigata by I. Bolivar. A
female of this material is at hand, bearing Bolivar’s identification label. The

|
|

F. WILLEMSE: The genus Cranae 139

specimen differs from the female of Form | als follows: elytra shorter, reaching
distal end of basal third of hind femur; subgenital plate flattened, not sulcate
apically; occipital spots not ovoid, but elongate; fore and middle femora reddish;
basal half of lower inner marginal area and fishbone pattern of outer medial area
of hind femur black.

Form 4
Cranae unistrigata: Kevan, 1966: 411 (Jobi I. only).

Material studied: Jobi Isl., Poue, T. Barbour (24, discoloured) (ANSP, MC).
Different from the holotype of unistrigata in the apical penis valves (figs. 28—29)
which are much shorter. Like in Form 2, the difference is difficult to evaluate.

Form 5

Material studied: Biak I., 5 m, 26.ix.1958, Pandanus, J. L. Gressitt (1 juv.g, 1 2)
(BPBM).

The phallic complex of the juvenile male is not yet sclerotized. The coloration
of the specimen is similar to that of the adult female. The latter differs from the
female of Form | in the coloration as follows: general colour of face and legs not
olivaceous green but yellowish-green; black fasciae of head and pronotum
narrower; distal part of hind femur not sanguineous red but suffused orange. This
form comes near the following one.

Form 6
Cranae unistrigata: Ramme, 1941: 90 (Farbungsrasse Sepik-Gebiet); C. Willemse, 1956: 107 (partim).

Material studied: Neth. Ind.—Amer. New Guinea Exp. 1938—39, Hollandia,
vii.1938, J. L. Toxopeus (29) & Bernhard Camp, 50 m, xii.1938, J. Olthof (19)
(NMM); Neth. New Guinea, Waris, S. of Hollandia, 450—500m, 1—7.viii.1959, T.
C. Maa (3g) (BPBM); Kais. Wilhelmsland, Paup, 1910, Dr. Schlaginhaufen (19)
(ZMHU); D.N. Guinea, Lager a. Töpferfluss, Kais. Augustafl. Exp., 29.iv.1912,
Burgers (19) & Standlager a. Aprilfl., 8—9.i.1912, Bürgers (13) (ZMHU); NE.
New Guinea, Adelbert Mts., Wanuma, 800—1000 m, 27.x.1958, J. L. Gressitt (14,
19) (BPBM).

This group of specimens from several localities in the northern part of central
New Guinea is uniform. It differs from Form 1 in the apex of phallus and in
coloration. The apical penis valves are slightly shorter, wider basally and more
tapering apically (figs. 30—31). The yellow occipital spots are conspicuously
elongate, the black fasciae of head and pronotum narrower, fore and middle legs
yellowish or olivaceous, hind femur yellowish or orange-yellowish and suffused
with orange or orange-red distally; femur more marked in the male than in the
female.

The material from Waris and Adelbert Mts. has not been recorded previously.

140 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

Cranae genjam sp.n.
(figs. 22—23, pl. 4 fig. 28)

Material studied: ¢ holotype, labelled: Neth., New Guinea, Genjam, 40 km W.
of Hollandia, 100—200 m, 1—10.iii.1960, T. C. Maa (BPBM).

Description.

Male, pl. 4 fig. 28. Differs from unistrigata as follows: face less wrinkled,
furculae larger, apical penis valves much shorter, wider basally and more strongly
tapering apically (figs. 22—23). Coloration much as in the holotype of unistrigata
except for the pronotum and hind femur. The black median fascia over pronotal
dorsum narrower, as wide as each of the yellow lateral parts of the dorsum. The
sanguineous distal part of hind femur much shorter, not extending over the inner
lower marginal and inner medial areas and better defined.

Female. Unknown.

Measurements (length in mm): body, 20.5; pronotum, 4.3; elytron, 9.1; hind
femur, 11.6.

Distribution: New Guinea (Genjam, near Hollandia).

Discussion. The species is well-defined by its resemblance to nominate
unistrigata in combination with its different apical penis valves. As to this
character, genjam is related with tibialis and patagiata.

Cranae trivittata C. Willemse, 1922
(pl. 2 fig. 12)

Cranae trivittata C. Willemse, 1922: 714, fig. 5; 1939: 75; F. Willemse, 1966a: 38; 1966b: 65 (Bivak Eiland
only).
Cranae trivittata trivittata: Ramme, 1941: 92; C. Willemse, 1956: 99, 105 (partim).

Material studied: © holotype, labelled: Z. Nieuw Guinea Lorentz 1909 - 10
Kloofbivak (ITZ).

Additional material: Z. Nieuw Guinea, Bivak Eiland, 1909—10, Lorentz (19)
(NMM).

Redescription.

Male. Unknown.

Female. Differs from the type-species as follows. Head comparatively larger and
rather more globose. Interocular distance wider. Fastigium verticis widely
triangular, more declivous, apex wider. Pronotum (pl. 2 fig. 12) shorter, sulci
weaker; dorsum more depressed, prozona, in profile, slightly upcurved; lateral
lobe shorter than high, posterior margin slightly concave. Elytron slightly longer,
reaching about middle of hind femur, apex narrower. Ovipositor valves more
slender.

Coloration as described in the original paper. Marked characters are: head pale
olivaceous green, with a black postocular fascia, yellow fascia along anterior
margin of eye and lower margin of cheek and a pair of yellow, elongate-ovoid

F. WILLEMSE: The genus Cranae 141

spots from between the eyes, almost reaching pronotum; pronotum yellow with
black fascia over upper part of lateral lobe and a median one over the dorsum, the
width of the latter almost equal to the width of either yellow, lateral part; elytron
black with yellow streak over the folding of elytron, at its base as wide as the
yellow lateral part of pronotal dorsum; fore and middle legs and hind femur pale
olivaceous green, the latter with fishbone pattern black, an ivory-white ante-
genicular ring, which is bordered by a narrow, complete, blackish ring proximally
(see original paper, fig. 5).

Measurements (length in mm): body, 26.0—27.0; pronotum, 4.5—5.0; elytron,
8.8—9.0; hind femur 14.7—15.1.

Distribution: SW New Guinea (Kloofbivak and Bivakeiland).

Discussion. The large head, short pronotum and coloration are distinct
characters. However, the male being unknown, the species cannot be properly
defined. The external morphology is much as in the unistrigata group. The
coloration resembles that of rufipes and manokwari.

Cranae rufipes Ramme, 1941
(figs. 32—33, pl. 2 fig. 13)

Cranae trivittata rufipes Ramme, 1941: 92; C. Willemse, 1956: 99, 106; 1962: 64.
Cranae trivittata trivittata: C. Willemse, 1956: 105 (partim) (?).

Material studied: 9 holotype, labelled: Deutsch Neu Guinea (ZMHU).

Additional material: NE New Guinea, Huon Peninsula, Finschhafen, 10 m,
16.iv.1963, J. Sedlacek (1 9) (BPBM); Nouv. Guinée, Baie Triton, 1841, Jacquinot
(19) (MNHN).

Redescription.

Male. Unknown (2), see below.

Female. Differs from the type-species in the slightly more globose head, wider
interocular distance, more widely triangular fastigium verticis with wider apex and
the slightly shorter elytra with narrower apex. Differs from trivittata in the less
globose head and longer pronotum (pl. 2 fig. 13), with the dorsum less depressed
and the prozona not upcurved.

Coloration differing from that in trivittata as follows: black median fascia over
pronotal dorsum slightly wider, nearly one-and-a-half times as wide as width of
either, yellow, lateral part of the dorsum; fore and middle femora reddish; hind
femur with antegenicular ring slightly wider and olivaceous green, bordered
proximally with a black ring, which is incomplete on the outer side.

Measurements (length in mm): body, 30.0-30.2; pronotum, 5.5—5.6; elytron,
7.5— 7.9; hind femur, 14.0— 14.9.

Distribution: NE New Guinea (Finschhafen and Triton Bay).

Discussion. Like trivittata, the taxon is not well-defined. The morphological
distinction between rufipes and trivittata is clear enough to raise rufipes to specific
rank. The Finschhafen female agrees completely with the holotype, while the
female from Triton Bay differs slightly in general colour, which is more orange-

142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

yellowish, and in the width of the antegenicular ring of hind femur, which is as,
narrow as in trivittata.

The following material is at hand: Kais. Wilhelmsland, Toricelli Gebirge, 1910, |
Dr. Schlaginhaufen (1 g, 1 9) (NMM); Neth. New Guinea, Waris, S. of
Hollandia, 450—500 m, 16—23.viii. 1959, T. C. Maa (2 9) (BPBM). The
morphology of this female material is very similar to that of the holotype of rufipes. 4
However, the coloration is slightly different: fore and middle femora olivaceous 4
yellow, black ring of hind femur complete, well-marked and as narrow as in |
trivittata, fishbone pattern of hind femur unicolorous with hind femur and yellow |
streak over elytron narrower. | |

By evidence of similarity, the pair from Toricelli Mts. is considered conspecific. |
The phallic complex of the male is much as in unistrigata, but the apical penis
valves are much wider (figs. 32—33). The two specimens were referred to
nominate trivittata by C. Willemse (1956). However, I am not certain whether this
material represents rufipes or a distinct taxon.

Cranae manokwari sp.n. |
(figs. 34—35, pl. 3 fig. 19, pl. 4 fig. 29) |
|
Material studied: & holo-, 9 allotype, labelled: New Guinea, Manokwari,
Tafelberg 150 m 22.11.63 R. Straatman (BPBM). |

Description. |

Male, pl. 4 fig. 29. Differs from the type-species as follows: head more globose,
interocular distance and fastigum verticis wider, elytron narrower with anterior
margin more evenly rounded; apical penis valves (figs. 34—35) rather tapering
apically, tips tooth-shaped and slightly outcurved.

Coloration as in rufipes, but vertex between the eyes and occiput en
black with two small, ovoid, yellow spots as in nominate unistrigata; median black
fascia of pronotal dorsum slightly wider and fore and middle femora olivaceous!
yellow. |

Female. Elytron, pl. 3 fig. 19. Abdominal terminalia as in type-species.
Coloration as in male, except upper side of head which is more yellowish- brown
with occipital spots faintly indicated.

Measurements (length in mm): body, & 22.4, 9 30.3; pronotum, ¢ 4.4, © 5.
elytron, & 7.8, 9 8.2; hind femur, g 11.9, 9 14.0.

Distribution: NW New Guinea (Tafelberg near Manokwari).

Discussion. The species is well-defined by the apex of phallus and the
coloration. It appears to be closely related to rufipes.

Cranae longipennis sp.n.
(figs. 14—15, 36—37, pl. 1 fig. 4, pl. 3 fig. 20, pl. 5 fig. 31)

Material studied: ¢ holotype, © allotype, labelled: Papua, Fly R. Olsabip 400°
—800 m, viii. 69, J. & M. Sedlacek (BPBM); paratypes: Neth. New Guinea Exp. | |
Star Range, Katem 200 m, 15.vi.1959 (23) (RNH).

F. WILLEMSE: The genus Cranae 143

Description.

Male, pl. 5 fig. 31. Differs from the type-species in body, head, tegmina,
abdominal terminalia and coloration. Body of larger size and more robust. Face
wider and more wrinkled. Fastigium verticis and interocular distance wider.
Tegmina longer, reaching end of middle third of hind femur. Elytron about three
times as long as wide and less tapering. Cerci more pointed and robust.

Phallic complex very large. Ancorae of epiphallus (figs. 14—15) partly joined
with the lateral plates. Apex of phallus (figs. 36—37) very robust, sheath of penis
conspicuously developed, tips of apical penis valves shell-shaped.

Coloration black, bluish-green, blue and yellow. Antennae blackish. Head
greenish-black with following parts cadmium-yellow: sulcus of fastigium verticis

44

Figs. 34—45. Cranae species, &, tips of apical penis valves and sheath of penis, ventro-posterior (even

numbers) and left lateral (odd numbers) view, similar scales; 34—35, manokwari sp.n. (holotype);

36—37, longipennis sp.n. (holotype); 38—39, rubra sp.n. (holotype); 40—41, kuekenthali Brunner
(paratype); 42—43, luctuosa C. Bolivar (Telaga, Obi I.); 44—45, pictipennis C. Willemse (allotype).

144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

and frontal ridge, scape, deep and symmetrical impressions of face, face along,
lateral keels and cheek along lower margin, the mouthparts partly, and a narrow |
stripe along dorso-posterior margin of eye. Pronotum yellow with three, longi- |
tudinal, wide, black fasciae: on each side, over the upper two-thirds of lateral lobe |
and a median one over the dorsum, being about twice as wide as the yellow lateral,
parts of the dorsum. Rest of thorax yellowish with episternum, upper part of

pleurae and margins and sutures of meso- and metasterna black. Elytron black ©

with wide, median, cadmium-yellow streak. Hind wing infumate. Abdomen
yellowish-brown, tergites laterally suffused green. Cerci and a median stripe on
supra-anal plate yellow. Fore and middle legs dark green with yellow streaks. Hind |

femur yellowish or olivaceous green, carinae and fishbone pattern dark olivaceous |
or bluish-green, inner lower marginal area completely dark blue. Hind knee dark |

blue with yellowish-white antegenicular ring, which is shorter than length of hind |
knee and incomplete ventrally. Hind tibia and tarsus bluish-black, tibia with a,
narrow, yellowish-white, postgenicular ring, hind tarsus yellowish laterally. |

Female. Larger than the male. Face, pl. | fig. 4. Elytron, pl. 3 fig. 20. DE
terminalia as in the type-species, but more robust. Coloration as in the male.

Measurements (lenght in mm): body, & 24.0—26.0, 9 33.2; pronotum, dg |

\
|
Il
|
|

4.6—5.0, © 6.3; elytron, g 11.3—11.6, 9 12.3; hind femur, G 12.5—12.7, 9 16.0.
Distribution: S. New Guinea (Katem, Star Range and Olsabip, Fly River).
Discussion. The species is very distinct. Shape and measurements of tegmina |

and phallic complex are conspicuous. The species seems related to the unistrigangi |

complex rather than to other species.

Cranae rubra sp.n.
(figs. 38—39, pl. 5 fig. 32)

Cranae patagiata; Krauss, 1903: 747, 759 (British New Guinea) (?)
Cranae sp. aff.patagiata; Kevan, 1966: 410 (not Normanby I.).

Material studied: ¢ holotype, 9 allotype, 9 paratype, labelled: New Guinea:

Papua Woodlark I (Murua) Kulumadau Hill, Mar. 19—22. 1957, W. W. Brandt, |

Cranae patagiata St.? det. D. K. mcE. Kevan, 1965 (BPBM).

Description.

Male. (pl. 5 fig. 32). Differs from the type-species as follows: pronotum narrower |
and shorter with dorsum more depressed and sulci slightly stronger; legs slightly |
more slender; apical penis valves (figs. 38—39) long, flattened laterally, tips

slightly pointed and recurved.

IE

Coloration black, carmine and yellow. Head and thorax as in unistrigata, but
occipital spots smaller. Elytron carmine, anterior margin widely, posterior margin
narrowly bordered with black. Abdomen and legs as in type-species, except forthe
hind femur. Hind femur in basal half carmine, in distal half blackish-red with a |

yellow antegenicular ring, which is almost twice as long as the length of hind knee.

Female. Larger than the male. Abdominal terminalia as in the type-species. |

Coloration as in the male, but face more unicolorous yellow.

F. WILLEMSE: The genus Cranae 145

Measurements (length in mm): body, ¢ 18.9, 9 23.6—24.5; pronotum, & 3.6, ©
4.5—4.8; elytron, & 8.2, 9 8.2—8.4; hind femur, Z 11.7, 9 13.0—13.5.

Distribution: New Guinea (Papua: Woodlark I.).

Discussion. The species is well-defined by the apex of phallus and the
coloration. Although the coloration is much as in patagiata, the species agrees
more closely with the unistrigata complex. The material studied by Krauss was not
at hand.

Cranae kuekenthali Brunner, 1898
(figs. 11—13, 40—41, pl. 3 fig. 21)

Ramme (1941) distinguished two subspecies, viz., kuekenthali and annulata. The
nominate form occurs in the northeastern part of Halmahera, annulata in the
western part of that island. Distinction was based on different coloration of the
hind femur. Ramme, in the same paper, synonymized the male of Cranae
marginata Brunner with nominate kuekenthali. Neither the type of marginata, nor
that of annulata or nominate kuekenthali could be studied (deposited in the
Senckenberg Museum Frankfurt). The only material available for the present
study are a paratypic male of nominate kuekenthali and some discoloured material,
labelled: Halmahera, T. Barbour (14, 2 9 ANSP, 1 & MC). As to the discoloured
material, agreement with kuekenthali is apparent, but subspecific distinction is
impossible. For the time being, no comments are given on Ramme’s opinion.

Cranae kuekenthali kuekenthali Brunner, 1898

Cranae kuekenthali Brunner, 1898: 198, 238, pl. 18 fig. 136 (partim); Kirby, 1910: 387; C. Bolivar, 1923:
146: C. Willemse, 1939: 74; Ramme, 1941: 88, 89; Kevan, 1966: 411.

Cranae kuekenthali kuekenthali; Ramme, 1941: 89; C. Willemse, 1956: 98, 102.

Cranae marginata Brunner, 1898: 198, 236, 237 (partim); Kirby, 1910: 387; C. Willemse, 1939: 74; Ram-
me, 1941: 87, 89 (¢ synonymy established); C. Willemse, 1956: 102 (in synonymy only).

Material studied: 14 paratype, labelled: Halmaheira 1894 W. Kükenthal leg.,
Paratypus (ZMHU).

Redescription.

Male. Differs from the type-species as follows. Integument of face and
pronotum smoother, scarcely pitted. Head larger; face less wrinkled, wider and
slightly depressed along lateral keels. Pronotum shorter, more cylindrical (dorsum
less depressed), in the middle slightly compressed laterally; sulci stronger; lateral
lobe not as long as high, posterior angle slightly pointed. Elytron shorter, reaching
about end of basal fourth of hind femur, about twice as long as wide, anterior
margin evenly and widely rounded, archedyction conspicuous. Furculae slightly
larger and closer together. Cercus more slender and extending slightly beyond tip
of subgenital plate. Ancorae of epiphallus not joined with lateral plates (fig. 11).
Cingular rami much wider (figs. 12—13). Apex of phallus shorter, roughly conical,
tip of penis valves excavated medially (figs. 40—41).

Coloration as given in the original paper. Marked characters are summarized as
follows: head yellow with wide, black, postocular fascia and narrower, black stripe

146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

from between eyes over occiput, widening posteriorly; pronotum black, margins
bordered with yellow, along anterior margin of dorsum widened into a pair of
yellow spots; elytron yellowish, apex and apical part of anterior margin bordered
with black; femora orange-red, the hind one apically more yellowish and with a
narrow, black ring placed just distally of middle of hind femur; hind knee blackish;
hind tibia bluish suffused with yellowish.

Female. Larger than male. Subgenital plate flattened, between the keels not
sulcate. Elytron as in pl. 3 fig. 21, in other specimens the black pigmentation may
extend more anteriorly. Coloration as in the male.

Measurements (length in mm): body, ¢ 21.0—22.0, 9 29.0—29.5; pronotum, &
4.1—4.4, Q 5.5—S.7; elytron, g 5.6—5.8, 9 5.2—5.3; hind femur, ¢ 12.4—12.6, 9
15.2—15.6.

Distribution: Moluccas (Halmahera).

Discussion. The species is well-defined by the external morphology, the phallic
complex and the coloration. The description of the female has been made after the
discoloured material recorded above (ANSP, MC). Whether the not sulcate
female subgenital plate is a reliable character is not certain, as the material is
shrivelled. A figure of the phallic complex is also given by Kevan (1966, Pacific
Insects 8: 697, fig. 2) under the name Cranae kuekenthali (nec I. Bolivar). The
species, together with luctuosa, forms a group (see below under /uctuosa).

Cranae kuekenthali annulata Ramme, 1941

Cranae kuekenthali Brunner, 1898: 198, 236, 238, pl. 18 fig. 36 (partim).
Cranae kuekenthali annulata Ramme, 1941: 89: C. Willemse, 1956: 99, 102; Kevan, 1966: 411.

No comments are given because of lack of material.

Cranae luctuosa C. Bolivar, 1923
(figs. 42—43, pl. 1 fig. 5, pl. 2 fig. 14, pl. 3 fig. 22, pl. 4 fig. 30)

Cranae luctuosa C. Bolivar, 1923: 145; C. Willemse, 1939: 74; 1956: 99, 103; F. Willemse, 1966b: 64.

Material studied: 1 ¢ paratype, labelled: Moluques Obi Major J. Waterstradt
1902 (NMM).

Additional material: Obi I., Telaga, 3 & 4 & 25 & 26.viii. 1953 (4 3, 1 9 NMM;
1 &, 1 9 RNH); Obil., Laiwui, 27.1x.1953 (1 9) (NMM).

Redescription.

Male, pl. 4 fig. 30. Differs from kuekenthali as follows. Head larger, more
globose. Pronotum, pl. 2 fig. 14; posterior margin of lateral lobe slightly concave.
Elytron longer, about three times as long as wide, anterior margin less widely
rounded. Supra-anal plate slightly wider and shorter, transverse ridges more
distinct. Cerci longer, extending well beyond tip of subgenital plate. Apex of
phallus slightly smaller and less conical (figs. 42—43).

Coloration as described by C. Willemse (1956). Marked characters are as

|
| F. WILLEMSE: The genus Cranae 147
|

| follows. Head from pale yellow to olivaceous yellow, with the following parts
| bluish-black: a transverse stripe over the face below the antennae, a wider one
‚along clypeal margin and lower margin of cheek, more or less the whole cheek, the

area behind the eye, a median stripe over the occiput and the posterior part of the
latter. Thorax bluish-black, margins of pronotum yellow or, usually, as spotted
| yellow: a median one on the anterior margin of the dorsum and the others on the

anterior and posterior angles of the lateral lobe. Elytron yellowish-white, apex and

apical part of anterior margin bordered with black. Fore and middle legs
| olivaceous green, more or less suffused with bluish. Hind femur in proximal two-
thirds bluish-black, in distal third yellow. Hind knee and tibia bluish-black.

Female. Larger than the male. Face, pl. 1 fig. 5. Elytron, pl. 3 fig. 22. Abdominal

terminalia as in kuekenthali. Coloration as in the male, but head, pronotum and
hind femur less black and occasionally almost completely yellowish.

Measurements as given by C. Willemse (1956).

Distribution: Moluccas (Obi I.).

Discussion. The species is well-defined by morphology and coloration. Its
resemblance to kuekenthali is striking. Both species are characterized, among
Cranae, by smoother integument, shorter and more cylindrical pronotum with
deeper sulci and another type of phallic complex. In these characters, the twe
agree with Opiptacris Walker rather than with Cranae. However, the developec
tegmina and the open tympanum do not agree with Opiptacris.

Cranae glabra sp.n.
(pl. 1 fig. 6, pl. 2 fig. 15, pl. 3 fig. 23, pl. 5 fig. 33)

Material studied: 9 holotype, 1 © paratype, labelled: Neth. New Guinea: Biak
I.: Kampong Landbouw, 50—100 m, May 27, 1959, J. L. Gressitt; 1 9 paratype:
Biak I. (Neth. N. Guinea) SE coast 5 m, xi—26—1958, Freycinetia, J. L. Gressitt; 1
juvenile 4 Neth. New Guinea; Biak I., Mokmer 5—10 m, v—26—1959, J. L.
Gressit (all BPBM).

Description.

Male. Cerci of the juvenile male long, extending well beyond tip of subgenital
plate. Phallic complex not sclerotized. Coloration pale yellowish.

Female, pl. 5 fig. 33. Differs from the type-species as follows. Body more
slender, integument much smoother. Face (pl. 1 fig. 6) not wrinkled and
comparatively narrow. Pronotum (pl. 2 fig. 15) shorter, more cylindrical, in the
middle slightly laterally compressed, sulci stronger, posterior angle of lateral lobe
slightly pointed and posterior margin slightly concave.

Elytron (pl. 3 fig. 23) smaller, much narrower, three times as long as wide,
reaching middle of second abdominal tergite, both elytra widely separated
dorsally; anterior margin slightly rounded, posterior margin straight, distally
tapering towards narrow, undivided, parabolic apex; veins rather reduced, not by
far reaching anterior margin. Hind wing almost as long as elytron. Tympanum as
usual. Legs slender, hind femur attenuate, fishbone pattern and keels moderately
developed. Subgenital plate flattened, not sulcated between low keels.

148 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

34

Plate 5. Figs. 31—34. Cranae species, lateral view: 31, longipennis sp.n., 3 (holotype); 32, rubra sp.n., 3
(holotype); 33, glabra sp.n., Q (holotype); 34, pictipennis C. Willemse, 3 (allotype).

F. WILLEMSE: The genus Cranae 149

Coloration yellowish and orange-brown. Antennae dark brown, apically paler.
Head yellowish, mouthparts mottled with bluish-green. Pronotum orange-brown
with three narrow, bluish-green fasciae: on each side over upper part of lateral
lobe and a median one over dorsum; these fasciae are widest along anterior and
posterior margins of pronotum, and almost obsolete near the middle of pronotum.
Pleurae with lower part orange-brown, upper part bluish-black. Pro-, meso- and
metasterna brownish. Elytron yellow, anterior margin broadly bordered with
black. Abdomen brown, proximal tergites blackish brown. Coxae orange-brown.
Fore and middle legs orange-brown, tibiae suffused with bluish-green. Hind femur
orange-brown, lower inner marginal area bluish, outer side basally slightly suffused
with dark brown. Hind knee, tibia and tarsus dark blue or bluish-green.

Measurements (9 length in mm): body, & 27.8—28.3; pronotum, 4.7—4.8;
elytron, 5.1—5.8; hind femur, 13.8—14.6.

Distribution: New Guinea (Biak I.).

Discussion. Although the adult male is not known, the species is well-defined.
Both morphology and coloration are quite distinct. Integument, head, pronotum
and female abdominal terminalia agree with kuekenthali and luctuosa. These
features agree with Opiptacris Walker rather than with Cranae. Besides, the
tegmina in glabra are much as in some species of Cranaella Ramme (compare F.
Willemse, 1977: 110). Apparently, Cranae glabra is more or less intermediate
between Cranae, Cranaella and Opiptacris.

Cranae pictipennis C. Willemse, 1932
(figs. 44—45, pl. 1 fig. 7, pl. 2 fig. 16, pl. 3 fig. 24, pl. 5 fig. 34)

Cranae pictipennis C. Willemse, 1932: 279; 1939: 74; F. Willemse, 1966a: 38; 1966b: 64; 1976: 121 (origi-
nal combination re-established).
Opiptacris pictipennis; Uvarov, 1937: 17, 18; C. Willemse, 1956: 95.

Material studied: type-series, labelled: Buru, Station Nal Besi (9 paratype,
RNH) & Station 16 (9 holotype, RNH) & Station 7 (¢ allotype, NMM), 1921, leg.
L. J. Toxopeus, Cranae pictipennis nov. sp. det. C. Willemse, Opiptacris pictipennis
Will. det. C. Willemse, type-labels. The specimens are discoloured.

Redescription.

Male, pl. 5 fig. 34. Differs from the type-species as follows. Body more robust.
Integument of face and pronotum (pl. 2 fig. 16) strongly pitted. Pronotum with sulci
less deep, posterior angle of lateral lobe obtusely pointed, posterior margin slightly
concave laterally, slightly emarginate dorsally. Tympanum closed, as a narrow,
hour-glass-shaped furrow. Elytron very short, reaching first abdominal tergite,
roughly circular with apex broadly rounded to almost truncated, venation
obsolete. Hind wing represented by a small membrane. Phallic complex as in type-
species, except for its apex. Sheath of penis not strongly folded apically but more
gradually merging into apical penis valves (figs. 44—45); the latter tapering
apically and excavated medially.

Coloration. The material is strongly discoloured. Elytra, apex of mandible,

150 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977 |

crescents of hind knee and furculae of the male, all black. A faintly retained |
pattern on the head and pronotum is much as in kuekenthali. |
Female. Larger than the male. Face, pl. 1 fig. 7. Elytron, pl. 3 fig. 24. Abdominal
terminalia about as in the type-species. Coloration as in the male. |
Measurements (length in mm): body, & 25.1, 9 28.1—32.0; pronotum, ¢ 4.7, A
5.3—5.3; elytron, ¢ 2.6, 9 2.8—3.0; hacken, 3 13.4, 9 14.8—15.1. |
Distribution: Moluccas (Buru). |
Discussion. The species is well-defined morphologically. In the tegmina and.
tympanum it agrees with Opiptacris Walker. However, the integument and |
especially the phallic complex disagree strongly with that genus. The integument is |
much as in Cranaella Ramme. As in the case of glabra and the kuekenthali-luctuosa . |
group, pictipennis forms a link between Cranae, Cranaella, and Opiptacris. |

'

Cranae emendata Brunner, 1898 |

|

Cranae emendata Brunner, 1898: 196, 236, 237; Kirby, 1910: 387; C. Willemse, 1939: 74; Ramme, 1941:
92; C. Willemse 1956: 98, 99 |

|

The species is known after a single, discoloured female from Borneo. The type.
is in the Senckenberg Museum, Frankfurt, and not at hand. As no further material |
of the genus is known from Borneo, the record certainly needs confirmation. |
|
The following five species are briefly described but pending the discovery of the.
male, no species name is proposed. |

Cranae sp.

Material studied: NE. New Guinea, Aseki, 19.xii.1965, H. Pyka (19) (author’ | |
collection). |

Near longipennis. Body robust, face wide and strongly wrinkled. Pronotum
rather depressed from above, tegmina wide and long, reaching beyond the middle
of hind femur. General colour black; yellow stripes over head and pronotum as in
unistrigata, but wider; elytron with a broad yellow streak; all femora red, the hind |
one with a yellow antegenicular ring and yellow streaks, one over the upper |
marginal areas and another along the lower outer carinula; hind knee black; all
tibiae blackish, suffused with red. |

Cranae sp.
Cranae sp. aff. patagiata; Kevan, 1966: 410 (Normanby I. only).

Material studied: Papua, Normanby I., Wakaiuna, Sewa Bay 1—10.x1.1956,

M. W. Brandt (2 juv. 3, 1 9) (BPBM).
Near trivittata, rufipes and manokwari. Head about as large as in trivittata, |
pronotum about as in rufipes. Phallic complex of juvenile males not yet sclerotized. :

|

F. WILLEMSE: The genus Cranae 151

Coloration as in rufipes, except for the legs. Fore and middle legs olivaceous. Hind
femur scarlet red with a broad, yellow antegenicular ring, bordered proximally by
a narrow black incomplete ring. Hind femur much as in rubra.

Cranae sp.

Material studied: NE New Guinea, Bubia, Markham R., 50 m, 20.ix.1955, J. L.
Gressitt (1 9) (BPBM).

Near manokwari but differing by more globose head and coloration of hind leg.
Hind femur olivaceous green, fishbone pattern bluish-black, antegenicular ring
orange-red, bordered proximally by a black dorsal spot. Hind knee and tibia
dark blue.

Cranae sp.
Cranae unistrigata; C. Willemse, 1956: 106 (Sabang only).

Material studied: Nieuw Guinea, Sabang vii.1907, Lorentz (1 9) (NMM).

Near unistrigata, but differing by more robust body, shorter pronotum and
coloration. Black fasciae of head and pronotum conspicuously narrower, median
one as wide as each of the yellow lateral parts of dorsum. Yellow streak over
elytron much wider. Hind femur yellow, fishbone pattern dark olivaceous, yellow
antegenicular ring narrower and bordered proximally by a narrower, well-
marked red ring.

Cranae sp.

Cranae unistrigata; Kevan, 1966: 411 (Roon I. only).

Material studied: D. N. Guinea, Roon Isl., Thomas Barbour (1 9) (ANSP). The
specimen is discoloured.

Near unistrigata, but rather more robust and head remarkably large and globose.
The discoloration being incomplete, the following pattern is still recognizable:
between the eyes a pair of black spots, behind the eye a wide black fascia,
pronotum unicolorous except for the central area of lateral lobe, which is darker.
Elytron as in unistrigata.

REFERENCES

Bolivar, C., 1923. Nuevas formas del grupo Cranae (Orth. Locustidae). — Boln R. Soc. esp. Hist. nat.
23: 145—150.

———, 1932. Estudio de un nuevo Acridido de Madagascar del grupe Cranae (Orth. Acrid). — Eos
Madr. 8: 391—396, figs.

Bolivar, I., 1898. Contributions à l’étude des Acridiens. Espèces de la faune Indo et Austro-Malaisien-
ne du Museo Civico di Storia naturale di Genova. — Ann. Mus. civ. Stor. Nat. Giacomo Doria
(2) 19: 66—101.

———, 1918. Estudios entomolögicos. Tercera parte. La sección Oxyae. — Trab. Mus. nac. Cienc. nat.
Madr., Zool. 34: 5—43.

Bruner, L., 1907. Subfam. Acridiinae. In: L. Bruner, A. P. Morse and R. Shelford, 1900—1909. Biol.
Centr. Amer., (Zool.) Insects, Orthoptera 2: 208—341.

152 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 5, 1977

Brunner von Wattenwyl, C., 1893. Revision du systeme des Orthoptéres et description des espèces rap
portées par M. Leonardo Fea de Birmanie. — Annali Mus. civ. Stor. Nat. Giacomo Doria (2)
13: 1—230, figs.

———, 1898. Orthopteren des Malayischen Archipels gesammelt von Prof. Dr. W. Kükenthal in den |
Jahren 1893 und 1894. — Abh. senckenb. naturforsch. Ges. 24: 193—288, figs. |

Haan, W. de, 1842—1844. Bijdragen tot de kennis der Orthoptera — Verh. nat. Gesch. Nederl. overz. |
Bezitt. 16 (Zool. 6): 45—124, pls. 11—20 (1842); 18 (Zool. 7): 125—164 (1842); 19—20 (Zool. |
8-—9): 165—228 (1843); 24 (Zool. 10): 229—248, pls. 21—23 (1844). |

Kevan, D. K. McE., 1966. Some Orthoptera—Caelifera from the Philippine, Bismarck and Solomon Is-
lands, with a few interesting records from New Guinea and the Molukkas. — Ent. Meddr. 34:
375—420, figs.

Kirby, W. F., 1910. A synonymic catalogue of Orthoptera. Vol. 3 Orthoptera Saltatoria. Part II. (Locus-
tidae vel Acridiidae). — London, pp.i—x, 1—674.

Krauss, H. A., 1903. Zool. Forschungsreizen Australien und Malayischen Archipel, 1891—1893 von Ri-
chard Semon. Bd. 5, Lief. 6. Orthopteren aus Australien und dem Malayischen Archipel ge-
sammelt von Professor Dr. Richard Semon. — Denkschr. med.-naturw. Ges. Jena 8: 745—770,
figs.

Ramme, W., 1941. Beitrage zur Kenntnis der Acrididen-Fauna des indomalayischen und benachbarter
Gebiete (Orth.). Mit besonderer Berücksichtigung der Tiergeographie von Celebes. — Mitt.
zool. Mus. Berl. 25: 1—243, figs.

Sjöstedt, Y., 1932. Orthopterentypen im Naturhistorischen Reichsmuseum zu Stockholm. 2. Acrididae. —
— Ark. Zool. 24A (1): 1—89, figs. |

Stal, C., 1873. Recensio Orthopterorum. 1. — Ofvers. K. Vetensk-Akad. Förh. Stockh., separ. pp.
1—20, 1—154.

———, 1878. Systema Acridiodeorum. — Bih. K. svenska Vetensk-Akad. Handl. 5 (4): 1— 100.

Uvarov, B. P., 1937. Some Acrididae from the Solomon Islands (Orthoptera). — Treubia 16: 15—20.

Walker, F., 1870. Catalogue of the specimens of Dermaptera Saltatoria in the collection of the British |
Museum. Part IV. — London, pp. 605—810. |

Willemse, C. (cf. Willemse, F., 1966, Bibliography of C. Willemse, Publties natuurh. Genoot. Limburg
16: 21—30), 1921a; 1922b; 1928b; 1932c; 1939c; 1956; 1962a. |

Willemse, F., 1966a. List of new taxa of Orthoptera, described by C. Willemse. — Publties natuurh. Ge- |
noot. Limburg 16: 31—42. |

———, 1966b. List of the types of Orthoptera in the collection of C. Willemse at the Natuurhistorisch _
Museum of Maastricht. — Publties natuurh. Genoot. Limburg 16: 43—73. |

———, 1972. A study of Pseudocranae I. Bolivar, 1898 and related genera Salinacris C. Willemse, 1956, |
Sphaerocranae gen. nov. and Malua Ramme, 1941 (Orthoptera, Acridoidea, Catantopinae). — _
Publties natuurh. Genoot. Limburg 22: 33—80, figs.

———, 1976. Studies on the Acridoid genera Opiptacris Walker and Bumacris Willemse (Orthoptera,
Acridoidea). — Tijdschr. Ent. 118: 117—158, figs. 1—44, 4 colourplates.

———, 1977. A study on the genus Cranaella Ramme (Orthoptera, Acridoidea, Catantopinae). — _
Tijdschr. Ent. 120: 109—120, figs. 1—30, pl. 1.

ROT

| DEEL 120

see MUS. COMP. ZOOL.
LIBRARY

AFLEVERING 6 MAR 8 1978 1977

HARVARD
UNIVERSITY

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

J. C. ROSKAM.

INHOUD

— Biosystematics of insects living in female birch catkins. I. Gall

midges of the genus Semudobia Kieffer (Diptera, Cecidomyiidae), p. 153—
197, text-figs. 1—122.

Tijdschrift voor Entomologie, deel 120, afl. 6 Gepubliceerd 30-XII-1977

BIOSYSTEMATICS OF INSECTS LIVING IN FEMALE BIRCH
CATKINS. I. GALL MIDGES OF THE GENUS SEMUDOBIA
KIEFFER (DIPTERA, CECIDOMYIIDAE)

by
J.C. ROSKAM

Division of Systematics and Evolutionary Biology, University of Leiden, The Netherlands
With 122 text-figures

ABSTRACT

Five species of Semudobia Kieffer have been studied in their developmental stages. All stages are
described and keys are provided. A neotype is designated for Cecidomyia betulae Winnertz, 1853, from
Betula pendula (fruit galls), The Netherlands, and four new species are described, viz., Semudobia
brevipalpis from Betula papyrifera (fruit galls), Canada, Quebec; S. steenisi from B. occidentalis (fruit
galls), U.S.A., Wyoming; S. tarda from B. pendula (fruit galls), The Netherlands; and S. skuhravae from
B. pendula (bract galls), The Netherlands.

INTRODUCTION

In this paper, the first of a series on the entomofauna of female birch catkins,
the galls, morphology and life cycle of five species of gall midges of the Holarctic
genus Semudobia Kieffer are studied. In a second publication, now in preparation,
morphology, life cycle and host relations of some of the hymenopterous parasites
of these midges will be discussed. Finally, a “food web” will be constructed and
attention will be paid to phylogenetic aspects of host-parasite relations of the
species under study.

The first description of Semudobia betulae was published by Winnertz in 1853.
He placed it in the genus Cecidomyia Meigen. Rubsaamen (1891) transferred the
species to his genus Hormomyia. Kieffer (1895) originally placed it in the genus
Oligotrophus Latreille, but in 1913 he made it the type-species of a new genus
Semudobia. Although a rather recent review was given by Barnes (1951), it is
necessary to discuss the literature in detail on certain topics. In fact these topics
anticipate-the splitting of the only described species in the genus.

Data on the number of antennal segments. — A mistake was made in the
original description by Winnertz (1853). Although there may be some variation in
the number of segments of the flagellum, the formulae 2 + 10/11 in the males (2
means scape plus pedicel, the other number is given for the segments of the
flagellum) and 2 + 10 in the females, as indicated by Winnertz, are highly
uncommon in Semudobia. Low (1878) critisized these numbers, but unfortunately
his counts (males 2 + 12, females 2 + 13) do not fit either, because males in

153

154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

Serıudobia usually have one segment more than females. Barnes (1951) had also
problems with the number of antennal segments, as for the females he recorded
2 + 12, “but occasionally (—) 2 + 15”, which must be an error for 2 + 13. The

data given by Rubsaamen (1891), Theobald (1892) and Kieffer (1913b) are correct: |

2 TA) 1118 (©).

Gall inducing and gall form. — Winnertz described C. betulae after specimens |
reared by Kaltenbach from female catkins of Betula alba L. from the preceding
year. Neither Winnertz (1853) nor Kaltenbach (1874) did note thatthe midge wasa

gall inducer. In fact, the statements by Kaltenbach (1874) and Binnie (1877) are
very confusing. They reported that the larvae live and pupate “in the interstices
between the scales’”, although Binnie also observed the “inside of a seed” as a
common position of the larva. Low (1878), and later Wachtl (1881), were the first
to study the behaviour of Semudobia carefully.

Low critisized Kaltenbach (1874) heavily, considering “diesbezügliche Be-
obachtungen als äusserst oberflachlich”, and observed the larvae of Semudobia
enclosed in galls. Löw and Wachtl both described the galls; they reported at least
two different gall forms but did not attribute these to different gall inducing
species. Rübsaamen (1891) added a third gall form to those described by Löw and
Wachtl, but did not suppose it to be due to a different gall inducing species either.
A remarkable mistake was made by Connold (1901, 1909), when he stated
Semudobia to be responsible for deformations of male birch catkins! These
deformations are not rare indeed and most causes were listed by Gäbler (1958).
The deformations of female catkins, attributed to Semudobia by Swanton (1912),
are doubtful too. Rübsaamen & Hedicke (1925— 1939) figured the three different
gall types (pl. XXV figs. 1—4) but again considered them to belong to one gall
midge species. The only authors, who realized the different gall forms in female
birch catkins to be due to different species, were Skuhravä & Skuhravy (1960,
1963).

Besides the gail inducing midges belonging to the genus Semudobia other gall
midges, inquilines, and phytosaprophagous midges may be found in the fruit
catkins of Betula. At least two, not identified, species of inquiline gall midges are
common in the galled catkins and may occasionally pupate “in the interstices
between the scales’’. So it seems probable that these inquilines were observed by
Kaltenbach (1874) and Binnie (1877). Suggestions about the identity of these
larvae were discussed later by Thomas (1893), Escherich (1942) and Skuhrava &
Skuhravy (1963). Hodges (1969) reported Clinodiplosis sp. as an inquiline midge.
Larvae of this genus are indeed frequent inhabitants of fruit catkins of Betula, but
there is no correlation between the occurrence of this midge and the presence of
Semudobia galls. Clinodiplosis is common in the ‘‘deformed and dwarfed catkins”
described by Swanton (1912) and is supposed to be a fungus eater in these catkins
(Skuhrava, 1970).

Larval characters. — Papillae of Semudobia larvae were studied by Rübsaamen
(1892), Kieffer (1895) and Mohn (1955). Phase-contrast microscopy being
unknown or uncommon in those days, it is not surprising that their data are very
incomplete and not suitable to distinguish the species. About the spathula

J.C. Roskam: The genus Semudobia 155

sternalis, the characteristic chitinized structure on the ventral side of the third
instar, more information can be found in the literature. Some variation is
apparent. Kieffer (1913b) described a wide structure: “spatule sessile, large, partie
basale presque transversale, partie antérieure moins large et bilobée”’. Rübsaamen
& Hedicke (1925—39) depicted a small and oblong one. Mohn (1955) depicted a
large and wide spathula. The description, given by Rübsaamen (1891), is
remarkable. He described a small, short, bilobed structure, “an dieser Platte
scheinen sich zwischen den beiden Lappen noch einige Zahnchen zu befinden”.
These small teeth were never reported again until Skuhrava & Skuhravy (1960)
depicted the spathula. They noted that this particular form of spathula only will be
found on larvae (Itonidae sp. in their terminology) living in galls that are
coalescent with the spindle of the fruit catkin. They also depicted the two spathula
types of Semudobia larvae living in fruit galls (in the strict sense), but supposed
these to be examples of intraspecific variation.

A short note on phenology. — Many authors indicated the end of March until
the beginning of May as the time of emergence of the adults. Forsius (1927), who
reported Semudobia from Finland, reared adults in August (probably under
unnatural conditions, as there are no receptive catkins at that time). This will be
discussed below (vide Phenology, pag. 180).

Other references of Semudobia. — Macquart (1853), Walker (1856), Dobner
(1858), Schiner (1864), Von Bergenstamm & Low (1876), Inchbald (1885), Lintner
(1887), Liebel (1889), Rubsaamen (1892), Von Tubeuf (1893), Collin (1904),
Houard (1908—13), Felt (1908, 1915, 1940), Kuster (1911), Ross & Hedicke (1927),
Docters van Leeuwen (1957), Gabler (1958), Buhr (1964), Mamaev & Krivosheina
(1965), Skuhrava & Skuhravy (1963), Bachmaier (1965), Gagné (1967).

Conclusion. — Differences in gall form (Rubsaamen & Hedicke, 1925—39) and
the larval spathula (Skuhrava & Skuhravy, 1960, 1963) have already been
recorded. These differences do not represent a range of intraspecific variation.
After a careful analysis of material from all over the northern hemisphere, these
differences are found to be supported by other characters concerning egg size,
papillary pattern in larval and pupal stages, number and form of antennal
segments, structure of genitalia and host parasite relations. Thus, it may be
concluded that they are differential characters, on which the genus Semudobia
may be divided into five species.

MORPHOLOGY

The gall (fig. 3).

Inflorescences of birches (Betula, Betulaceae) are monoecious. The female
birch catkin (fig. 1) consists of an elongated spindle bearing spirally arranged
condensed dichasia (Porter, 1967). In each dichasium a leaf-like scale bears three
axillary female flowers; the fruits are winged nutlets. In this paper, for
convenience’ sake, the scale is called “‘bract’’. This is not quite correct, because in
the morphological sense it is a combination of a primary and two secondary bracts
(fig. 2). Galling of tissues in the female catkin is exclusively the result of Semudobia

156 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

Figs. 1—8. Life cycle of Semudobia. 1, fruit catkin of Betula; 2, scale with female flowers; 3, scale with

fruits, the middle one infected; 4, egg; 5, larva; 6, pupa; 7, male; 8, female. Al—A8, abdominal

segments; AS, anal segment; H, head; SS, supernumerary segment; TH, first thoracic segment; ap,

apical papilla; as, apical spine; cd, dorsal collar papilla; cv, ventral collar papilla; d, dorsal papilla; |,

lateral papilla; Ifp, lateral facial papilla; Is, lateral spine; p, pleural papilla; pb, primary bract; ph,

prothoracic horn; s, sternal papilla; sb, secondary bract; t, terminal papilla; v, ventral papilla. 3, x 7; 4,
x 100; 5,6, x 35; 7,8, x 25. 1,2, after Strassburger et al., 1971.

activities. Semudobia galls are never observed outside female birch catkins. Galls
may be found when infected catkins are sifted out by separating bracts and nutlets.

Eggs are deposited between the bracts and flowers. Gall forming is induced by
the newly hatched larvae, which select the tissue to be infected. The different
species prefer different tissues and this results in different gall forms. Detailed
information about these forms will be given in the descriptions of the species.

As a rule, only one individual infects one bract or one ovary. Sometimes
however, several (up to five!) individuals, which may belong to different species,
are found in one single fruitlet. When not attacked by other insects, all these
larvae will develop into normal adults. About at the beginning of the second
instar a characteristic window-like spot (fig. 97) in the galled fruit is formed by

J.C. Roskam: The genus Semudobia 157

A

PE Coe

Fig. 9. Egg of S. tarda, detail of micropyle; 10—11, anal segment of first instar larva: 10, S. betulae, 11,
S. skuhravae; 12—13, pupa of S. skuhravae: 12, dorsal aspect of thorax, 13, anterior ventral papillae on
7th abdominal segment; 14, S. betulae, 3, head; 15—16, g, antennal scape: 15, S. betulae, 16, S.
brevipalpis; 17, thorax of S. betulae, &, lateral aspect; 18—21, wing of S. betulae, 3; 18, venation, 19,
detail of base, 20, sensory-pores on distal part of RI, 21, detail of tip. a, arculus; d, dorsal papilla; mm,
mesanepimeron; ms, mesanepisternum; ph, prothoracic horn; s, stigma; sp, sensory-pore; t, terminal
papilla. 9—11, x 325; 12, x 40; 13, x 250; 14, x 105; 15—16, 19—21, x 165; 17—18, x 35.

some of the species. This window functions as a “weak spot”, through which the
adults emerge. Some authors (Kieffer, 1895; Pitcher, 1957 and Hodges, 1969)
supposed the window to be made by the larval spathula. This can not be correct in
this case, because the spathula is only present in the third instar larva.

The egg (fig. 4).

Eggs may be found in the flowering female catkins of birch; they are scattered
between the bracts and flowers. For detailed information about oviposition, see
Hodges (1969). Freshly laid eggs are transparent, becoming orange-reddish when
the larva develops. Semudobia eggs are centrolecital, the chorion is smooth and
one micropyle is visible on the side where in the ripe egg the anal segment of the
embryo will be situated (fig. 9). Egg sizes are taken of ovarian eggs and statistically
more or less different egg sizes are found for the individual species (fig. 22). Per

158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

220 240 | 260 280 300 380
egg length (A)

|
|
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|
|
|
320 340 so
|
|
|
|
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egg length (A)

Fig. 22. Egg lengths of Semudobia. a—d, histograms: a, S. skuhravae, b, S. betulae, c, S. brevipalpis, d, S.
tarda; e, corresponding frequency distributions. na, mean value of S. skuhravae, nb of S. betulae, hc
of S. brevipalpis, nd of S. tarda (S. steenisi not studied).

J.C. ROSKAM: The genus Semudobia 159

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antennae (b) for each of the three larval instars of S. betulae.

=

female five eggs are measured; four females are taken per population and three
populations per species.

The larva (fig. 5).

Characters of all larval stages are of great value in gall midge systematics on the
generic level. Moreover, they prove to be essential for the delimitation of the
species of Semudobia. Semudobia midges pass through three larval stages. In all
instars the larva consists of a weakly sclerotized head (H), a supernumerary
segment (SS), three thoracic segments (Th 1—3) and nine abdominal segments, the
last of which is the aral one (A 1—8, AS). From the dorsal side of the head-
capsule arises a pair of short antennae. On the ventral side of the antennae a small,
oval area is visible. The length of the antenna does not differ remarkably among
the different species; it is 2 u in the first instar, 6 u in the second and ll yp in the
third. Eye-spots are distinct in all stages, but not in the last inactive period of the
third instar in which diapausis is passed. In this full-grown larva the head is
retracted into the supernumerary segment. Measurements of the head-capsule are
given in the descriptions. As demonstrated in fig. 23, there is no correlation

160 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

between length and width of the capsule in any instar. An interstadial growth (as
was found by Den Hollander, 1975, for Tipula) does not occur in larvae of these
midges. Semudobia larvae are apodous. The colour of the first instar larva is pale
orange, with a bright red spot where the mid gut is situated. There are 9—10 pairs
of caeca. The second instar larva is yellowish pale orange and the third instar
bright orange; when full-grown, it has a reddish tinged thorax.

Respiration in the first instar is, depending on the species, apneustic or
metapneustic; when stigmata are developed, they are situated on A 8. In second
and third instar larvae respiration is peripneustic; nine pairs of stigmata are
developed on Th I and A 1—8.

Papillae are developed in a very definite pattern, that is the same for all instars
but different for the various species. The terminology of Mohn (1955) is followed.
Papillae are very distinct in the first larval stage. In that stage, only the dorsal,
pleural and lateral papillae of A 8 and the terminal papillae of AS have a short seta
(up to 1 u). In the second and third instars, however, usually all papillae have a
short seta (— 2 u). The papillary pattern of the head appears to be very complex
and is not dealt with in this study. Depending on the species, the supernumerary
segment may bear one pair of dorsal collar papillae (CD) and/or one or two pairs
of ventral collar papillae (CV). On the dorsal surface of the thoracic segments, one
or two pairs of rows of dorsal papillae (D) are developed, laterally there are two
pairs of rows of pleural (P) and one or two pairs of rows of lateral papillae (L,
subventral in position). On the ventral surface of Th 1 one pair of sternal papillae
(S) is always distinct. The segments Th 2, 3 possess one pair of ventral papillae (V).

L THORAX ABDOMEN
YA AS

Fig. 24. Diagram of papillary pattern and tracheal system of third instar larvae, lateral aspect. a, S. tarda;

S. betulae; c, S. brevipalpis; d, S. steenisi; e, S. skuhravae. Al—8, abdominal segments; AS, anal

segment; CD, dorsal collar papilla; CV, ventral collar papilla; D, dorsal papilla; H, head; L, lateral

papilla; P, pleural papilla; S, sternal papilla; SS, supernumerary segment; T, terminal papilla; THI—3,
thoracic segments; V, ventral papilla; O, stigma.

J.C. ROSKAM: The genus Semudobia 161

The dorsal surface of the abdominal segments A 1—7 offers the best diagnostic
characters in papillary pattern. Depending on the species, 1—3 pairs of rows of
dorsal papillae may be developed. Laterally one pair of rows of pleural papillae
and one pair of rows of laterals can be observed. On the ventral surface of the
segments A 1—7 there may be one pair of anterior papillae and/or one pair of
posterior papillae. The anterior ventral papillae are in most species only visible in
the young first instar. The A 8 segment has a more simple pattern. One pair of
dorsal papillae is present in all species except one, in another species there are
usually no lateral papillae. In general, one pair of both pleural and ventral papillae
are present. Depending on the species, 1—3 pairs of papillae are present on the
anal segment. In second and third instar larvae papillae are situated outside the
anal field (a distinct area around the anus); these may be distinguished as terminal
papillae (t, fig. 85). In the first instar larva, where the delimitation of the anal field
is not clear (fig. 10, 11), the discrimination between anal and terminal papillae may
be a moot point. Contrary to Mohn (1955), Hodges (1969) considered a papilla a
simple skin-structure and not a sense-organ. The statements by Kieffer (1895)
and Mohn (1955) that pleural papillae are not developed in Semudobia must be
imputed to the less sophisticated microscopy techniques of that time. Papillary
patterns are scheduled in fig. 24.

Skin structures, such as well developed spinule rows, are visible on the ventral
and dorsal surfaces of the first instar larvae. On the dorsal surface of second instar
larvae spinule rows may be less distinct. In regard to spinule rows it is possible to
divide larval segments in an anterior and a posterior part. In the anterior part, the
spinule rows are always distinct and they lie close to each other; in the posterior
part, however, they lie further apart or, more often, are absent. The spinule rows
are counted and the numbers are given in the descriptions. The first number
concerns the posterior part of Th 3, the second relates to the anterior part of A 1.
Between brackets the variation in number is given usually for ten specimens. Skin
structures and papillae are distinct in newly hatched first instar or just moulted
second instar larvae, but they become less distinct in older first and second stages.
The third instar larva has a verrucose surface. Dorsally there are no remarkable
differentiations, but ventrally the verrucae unite to form striae, while laterally they
tend to be isolated and more rounded. In the figs. 72, 73, e.g., small fields are
indicated that have no verrucose sculpture. On the ventral surface of the third
instar prothorax (Th 1) lies a heavily chitinized structure: the spathula sternalis.
Shape and size of the spathula, as suggested earlier, give good characters for the
delimitation of the species.

When full-grown (about mid August, in The Netherlands), the larva produces a
whitish membraneous cocoon (see also Hodges, 1969) and is ready to hibernate.

The pupa (fig. 6).

In morphology and descriptions of the pupal stage mainly Mohn (1961) is
followed. The colour of the pupa is bright orange. On the head, near the place
where the adult pedicel will develop, two heavily chitinized blunt apical spines are
formed. These spines help the pupa to perforate the window-pit of the gall just

162 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

before the adult emerges. Next to the apical spines two subrotund laterals are |
visible. Frontal spines are absent, on the ventro-lateral part of the future adult eye, .
one or two pairs of papillae are situated. Just caudal of the apical spines, a pair of

slender apical papillae is present. From the dorsal side of the prothorax a pair of

stout prothoracic horns (fig. 12) arises, on the end of which the prothoracic ©

|
|

stigmata are located. The papillary pattern on the abdominal segments A 1—8 is in

conformance with those of the larval stages and here also offer the best diagnostic
characters. In the pupa, the anterior ventral papillae are often distinct (fig. 13). On
the dorsal surface of A 1—7 a pair of rows of sensory-pores is also formed (fig. =:
The abdomen is covered with tiny, pointed setulae (fig. 13).

The adult.

In the descriptions of the adult stage mainly Harris (1966, 1968) is followed. For
further details, see Gagné (1968, 1973) and Yukawa (1971), from whom some of
the terms used are borrowed.

Male (fig. 7). — Body reddish-orange, with dark brown sclerotized parts. Eyes
holoptic (fig. 14), with a slight, incomplete, median incision. Antenna with scape,
pedicel and, depending on the species, 12—14 flagellomeres. The distal flagel-
lomeres are often fused. If the last flagellomere is more than 1% times the length
of the penultimate one, the total count of flagellomeres is given as ‘“‘n*” in the
descriptions, rather than ‘n’. Flagellomeres uninodal with subquadrate internodes
(stalks); sensorial spines present, usually two per flagellomere and situated on
antero-distal portion (fig. 33); circumfila simple, not looped; basal setae on horse-
shoe-shaped sockets. Scape larger than pedicel, with a ventral area with very short
setae and laterally and distally fields with long setae (fig. 15). Maxillary palpi three
segmented.

Numbers of setae on mesanepisternum and mesanepimeron (fig. 17) are given in
the descriptions. Sc of wing usually incomplete or absent, and indicated only by a
number of sensory-pores (fig. 19); R5 well separated from RI, and joining C near
wing apex. Cu halfway forked in Cul and Cu2; m 3 + 4, Cul and Cu2 only
indicated as wing-folds; Rs absent. Tarsal claws simple, bent nearly at right-angle
and a little shorter than empodia.

Abdominal terga 2—7 not strongly sclerotized, caudal row of setae present,
basal row absent, only one pair of short setae cephalad of caudal row; sterna 2—7
with both basal and caudal rows of setae, one pair of median short setae cephalad
of the basal row. Basimere (fig. 46) of genitalia stout with in internal angle a large
claspette (mesolobe); distimere stout, in some species a little inflated, about half as
long as basimere and with a dense row of strong teeth distally; superior lamella
(tergum 10) with V-shaped incision, forming a pair of lobes which are rounded
distally; inferior lamella (sternum 10) narrower than superior lamella and,
depending on the species, shallowly emarginated to with V-shaped incision;
aedeagus rather thick, distinctly shorter to a little longer than claspette.

Female (fig. 8). — Antenna with scape, pedicel and 11—13 subsessile
flagellomeres. Ovipositor retractile, tapering to a simple upper lamella and a short
to very short lower lamella (fig. 56, 57).

J.C. ROSKAM: The genus Semudobia 163

METHODS

Measurements of galls are taken in dry condition; in the case of fruit galls
stigmata and wings are excluded. Sizes of galls given are only valid for the host
plant species mentioned in the descriptions and for the galls inhabited by one gall
midge only.

Eggs are dissected from identified females and macerated in warm (85-90°C)
80% lactic acid. After being straightened under a cover-slip, length and width are
measured in the same medium. Midges, from which ovarian eggs are taken, are
preserved on a 70: 15: 15% aethanol-glycerol-water mixture. Eggs of females
preserved in other ways may not give comparable results.

Preparation of the larvae, especially of first and second stages, may cause many
difficulties. Therefore rather detailed information is given here about techniques.
Newly hatched larvae are collected from catkins just after flowering. These
catkins are freshly deep-frozen or preserved on a 70 : 15 : 15% mixture of
aethanol-glycerol-water. Older larvae are dissected from fresh or dried (herbarium
specimens) galls and preserved either dry, or in the same aethanol-glycerol-water
mixture. Fresh larvae are macerated in warm 80% lactic acid. Larvae from dried
galls first have to be hydrated carefully in a series of 90, 70, 30% solutions of
aethanol and in water before maceration in 80% lactic acid is possible. Swelling of
the body content, which might result in total distortion of the larval skin, is thus
prevented. Small larvae, first instar and young second instar, are examined in the
maceration medium. For a slide collection these larvae are mounted in polyvinyl-
lactophenol (Chroma Gesellschaft Schmid & Co., Stuttgart, Germany). To avoid
shrinking, the very tender skins are transferred to the mounting medium without
neutralizing or rinsing. The older larvae are washed in water after maceration and
the residuals of body contents are very carefully removed. The macerated larvae
are stored and examined in glycerol.

Pupae are dissected from the galls and macerated in 80% lactic acid. After
rinsing the pupae have to pass a series of 30, 70, 96% aethanol before being
mounted in euparal.

Measurements of all immature stages are taken from specimens straightened
under a cover-slip.

To rear the adults, galls are sorted according to the identification key for the
galls on page 166. The samples, mixed with peat-litter and deposited in glass-vials,
are allowed to pass diapause under more or less natural conditions. From January
on, the glass vials are transmitted to sand-filled flowerpots and kept in a climate
chamber (20°C, long day light regime). The cultures are moistened with 1%
propionic acid to prevent damage by mould infections. After emergence the adults
are promptly killed in a mixture of 70 : 15 : 15% aethanol-glycerol-water and stored
in the same medium. Because maceration might disturb tiny organs as antennae,
the specimens are directly dehydrated in a series of 30, 70, 96% aethanol. After
clearing in oil of cloves the specimens are mounted in euparal. On the slide the
head, left wing and male genitalia are mounted under separate cover-slips. The
body is slide-mounted in a lateral position, the head with the face up, the genitalia
dorsal surface up.

164 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

For examination a phase-contrast microscope (Zeiss, objectives Ph 10, 25, 40 |

and 63 x) is used.

TAXONOMIC PART

Semudobia Kieffer

Cecidomyia Meigen (partim); Winnertz, 1853: 234.
Hormomyia Rübsaamen, 1891: 137.

Oligotrophus Latreille (partim); Kieffer, 1900: 22.
Semudobia Kieffer, 1913a: 55.

Type-species: Cecidomyia betulae Winnertz, 1853:234.

Keys to species of Semudobia

(Egg sizes are not used in the keys because their accurate analysis was only
performed in the ovarian stage; fig. 22.)

Males
I. Third palp segment shorter than the second; claspettes rectangular, distally
emarginated; mainly/Palaearctiespecies sent 2.200 tarda sp. n.
— Third palp segment longer than the second; claspettes more or less triangular,
never distally emarginated)) UR eo ea EE 2
2. Ultimate flagellomere rather rounded, internodes transverse to subquadrate;
first and second palpal segments not broadly fused; claspettes verrucose;
aedeagus usually a little shorter than claspettes ........ skuhravae sp.n.
— Ultimate flagellomere rather pointed, internodes oblong; first and second
palpal segment broadly fused; surface of claspettes only finely sculptured;
aedeagus always a little longer than claspettes .................. 3
3. Antenna with 14 flagellomeres, at least 3 times the width of the head;
internode of third flagellomere at least 0.5 times the length of the node of this

segment West-Nearctieispeciestnn RO cen ONE ere steenisisp.n.
— Number of flagellomeres less than 14, length of antenna never more than 2.6
timessthe width'ofithe head mms 3 PE RE 4

4. Never more than one seta on distal surface of scape and then laterally placed
(fig. 16); third palpal segment at least 1.7 times the length of the second; at
least 5 setae on mesanepisternum and 8 on mesanepimeron; inferior lamella
about half-way with U-shaped incision, lobes acute; East-Nearctic species . .

EEE ee ners UN ER HELD ee Tee. Sik. Sees brevipalpisispaur

— 3—10 setae on distal surface of scape (fig. 15); length of third palpal segment
not more than 1.4 times the length of the second; not more than 3 setae on
mesanepisternum and not more than 5 on mesanepimeron; inferior lamella not
more than for about one-third V-shapedly incised, lobes rounded; mainly
Palaearctiesspecies ul UNE SiN Dr... es a betulae Wtz.

=

1.

DI

J.C. RoskAM: The genus Semudobia 165

Females
Tergum 8 triangular, length of this sclerite about equal to its width; ovipositor
always longer than hind femur, with heavily chitinized vaginal furca; inferior
lamella very short and truncate; mainly Palaearctic species ... tardasp.n.
Tergum 8 sub-triangular or rectangular to H-shaped, always somewhat longer
than wide; ovipositor shorter than hind femur; vaginal furca not remarkably
chitinized; inferior lamella at least a little shorter than wide, rounded ... 2
Ultimate flagellomere rounded; first and second palpal segments never broadly
USERS IRE NEN: skuhravae sp.n.
Ultimate flagellomere pointed; first and second palpal segments broadly fused

Antenna at least two times the width of the head; West-Nearctic species
NN: ie Sr ciel RIS TREE LEER END AS NH ei steenisi sp. n.
Antenna never more than two times the width of the head .......... 4
Distal setae on scape, when present, then placed laterally; third palpal segment
at least 1.5 times the length of the second, length of tergum 8 never more than
1.5 times its width; length of superior lamella about 1.3 times its height; East-
INGANG ro OE ee CR brevipalpis sp. n.
Distal setae on scape, when present, never placed laterally; third palpal
segment never more than 1.5 times the length of the second; length of tergum 8
about two times its width; length of superior lamella at least two times its
heichtzmamlyBalacarnctigspeciess 2 2 ee betulae Wtz.

First instar larvae!)
(No stigmata, or only one pair of stigmata developed, spathula sternalis absent.)
No stigmata developed at all; only one pair of rows of dorsal papillae on
abdominal segments A 1—7, no dorsal papillae on A 8 (the penultimate body

SCLC M ET A Een EN ee Nesey: en tae tarda sp. n.
One pair of stigmata on A 8; at least two pairs of rows of dorsal papillae on
scpments Alone DARON ANSE iS 0 fe e 2

Three pairs of rows of dorsal papillae on A 1—7, one pair of dorsal collar
papillae, on A 8 pleural and lateral papillae both developed skuhravae sp. n.
Two pairs of rows of dorsal papillae on A 1—7, no dorsal collar papillae, on A
Sinojlateralipapillaeidevelopedie PER RR RTE betulae Witz.

Second instar larvae
(Stigmata developed on first thoracic segment and first to eighth abdominal

segments; no spathula sternalis. Same key as for third instar, characters based on
spathula excluded.)

)

Nearctic species not studied and therefore excluded from this key.

166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

Third instar larvae
(Nine pairs of stigmata (vide second instar) and spathula sternalis developed.)
Spathula sternalis with a variable number of smaller teeth (as if broken off)
between the usually larger lobes; three pairs of rows of dorsal papillae on A
ESTER IND EEE DIET DEN INNERN IP skuhravae sp. n.
Spathula aiways bilobed; only one or two pairs of rows of dorsal papillae on A
1-7: Balmopn bis eds Sa Ders Die ENTE 2
Spathula large, lobes at least 15 u high; only one pair of rows of dorsal papillae
OTA N es Re RN OENE EN tarda sp. n.
Spathula smaller, height of lobes never more than 10 u; always two pairs of
rows of dorsal.papillae . 4.2 … vaten oe 2. nn i LIRE 3
Spathula oblong, at least 50 u high; no lateral papillae on A 8, only one pair of
terminal papillae; mainly Palaearctic species ........... betulae Wtz.
Spathula subquadrate to transverse, never more than 50 u high; lateral papillae
on A 8 present, 2—3 pairs of terminal papillae nnn 4

Lobes of spathula at least 5 u high, one pair of rows of lateral papillae on |

thoracic segments, two pairs of rows of ventrals on segments A 1—7; West-

Neasctieispecies. .. ., ech ne op ce can cues III Steenisisp.n. |
Lobes of spathula never more than 5 u high, two pairs of rows of lateral |
papillae on thoracic segments, one pair of rows of ventrals on segments A |

li East-Nearcticispecies: ct ee ee ree eee brevipalpis sp. n.
Pupae!)

. Two pairs of rows of dorsal papillae, dorsally of sensory-pores, on abdominal
segments Al nn 0 ues eee A RE skuhravae sp. n.
Only one pair of rows of dorsal papillae, dorsally of sensory-pores, on A 1—7

EERE ear a can che Mii ELI MS Oa MEA A PI 2

. No dorsal papillae laterally of sensory-pores on A 1—7, dorsal papillae on A 8
ADS fe Mera tee ilar BAT EE NEEN tarda sp. n.
One row of dorsal papillae laterally of sensory-pores on A 1—7, one pair of
dorsal papillacion' A 8c AE et eee 3
Prothoracic horn longer than 120 u, about 0.08 times the body-length or more;
mainlyBalacarcticispeciesi.n er. me ee n betulae Wtz.
Prothoracic horn shorter than 100 u, about 0.06 times the body-length or less;
West-Nearetiespecies 22.0... 20. DE EEN steenisi sp. n.

Galls
Either the axil of the scale is galled, or the galled fruit is distinctly coalescent
Withithe/scaleswindow-pitabse teen skuhravae sp. n.
Galling of the fruit, the gall is in ripe situation never coalescent with the scale
EE à 6 o o o 0 0 0 = 2

. Window-pit absent or indistinct, gall subrotund and glabrous, wings of fruit
oftentcompletely reduced 20 000 yn ee tarda sp. n.

') S. brevipalpis not studied and therefore excluded from this key.

J.C. ROSKAM: The genus Semudobia 167

= 3 Window=pitfs);.disuineti no er RIE ee ROSE cee Beaten 3
3. Often a window-pit on both sides of the gall, one of them large; gall glabrous,
wings of fruit often completely reduced; East-Nearctic species brevipalpis sp. n.
— Only one window-pit developed; gall more or less hairy, wings present .. 4
4. Gall usually concave on ad-axial side, window-pit irregularely formed; West-
INCARCHESPECIE Sa eh er fees ORI repens Le I ale steenisi sp. n.
— Gall usually convex on both sides, window-pit subrotund; mainly Palaearctic
SPECICSERE EN II IA ERRE. betulae Wtz.

Semudobia betulae (Winnertz)

(male: figs. 14, 15, 17—21, 32, 33, 42, 43; female: figs. 40, 41, 50, 54, 55, 102; imm. stages: figs. 10, 64, 65,

70—75, 95—97)

Male. — Width of head 1.17 (1.12—1.30) times its height. Antenna (figs. 32, 33)
with 2 + 12 segments, its length 2.53 (2.49—2.60) times the width of head; scape
ventrally with 2 (2—4) very short, 4 (3—4) lateral and 10 (3—10) dorsal long setae;
third flagellomere with its node 1.07 (1.00—1.08) times as long as its diameter;
internodes oblong, internode of third flagellomere 0.48 (0.38—0.48) times as long
as node. Maxillary palp 0.24 (0.24—0.28) times the width of head; third segment
1.27 (1.14—1.38) times as long as the second; first and second segments partially
grown together. Fronto-clypeal setae 7 (2—7); 2 (2—3) setae on mesanepisternum,
5 (4—6) on mesanepimeron. Wing length 1.68 (1.56—1.74) mm, 2.0 (1.9—2.2)
times as long as wide; RS 2.3 (2.2—2.3) times as long as RI; Sc indicated by 4
(3—4) sensory-pores, 4 (3—5) sensory-pores on basal part of Rl and 2 on distal
part, 3 pores on medio-distal portion of RS. Tibia of hind leg 0.94 (0.78—0.98) times
as long as femur, second tarsal segment 0.54 (0.46—0.57) times this length; fifth
tarsal segment 0.85 (0.83—0.85) times as long as the fourth. Distimere (fig. 42) 0.47
(0.47—0.58) times as long as basimere; inferior lamella weakly emarginate to
incised for about one-third, the incision broadly V-shaped, its sides converging to
its apex (fig. 43); aedeagus a little longer than the usually weakly chitinized,
rounded triangular claspettes.

Female. — Width of head 1.15 (1.15—1.28) times its height. Antenna (figs. 40,
41) with 2 + 11+ (10+—12) segments, its length 2.00 (1.44—2.00) times the width
of head; scape ventrally with 3 (2—3) very short, 3 (2—3) lateral and 3 (0—3)
dorsal long setae; third flagellomere with its node 1.08 (1.04—1.09) times as long as
its diameter. Maxillary palp (fig. 102) 0.28 (0.24—0.30) times the width of the head;
length of third segment 1.46 (1.25—1.46) times the length of the second; first and
second segments partially grown together. Fronto-clypeal setae 10 (1—10); 3
(2—3) setae on mesanepisternum, 7 (5—7) setae on mesanepimeron. Wing length
2.00 (1.66—2.14) mm, 2.1 (2.0—2.1) times as long as wide; RS 2.4 (2.3—2.4) times
as long as RI; Sc indicated by 5 sensory-pores, 6 (4—6) sensory-pores on basal part
of RI and 2 on distal part, 3 on medio-distal portion of RS. Tibia of hind leg 0.89
(0.88—0.90) times as long as femur, second tarsal segment 0.43 (0.41—0.46) times
this length; fifth tarsal segment 0.86 (0.83—0.92) times as long as the fourth.
Tergum 8 subtriangular (fig. 50), the height of this sclerite about 2 times its width.
Ovipositor (fig. 55) 0.75 (0.75—0.87) times as long as hind femur, for one- to about

168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

three-quarters retracted into the eighth abdominal segment; vaginal furca not
remarkably chitinized; superior lamella 2.5 (2.3—3.0) times as long as high;
inferior lamella with a length about equal to its width (fig. 54).

Immature stages. — Egg length n 326 u, n + 2 SD. (95% of egg lengths) between
312—340 u; width n 106 u. First instar larva (figs. 64, 65); length 255 (221—642),
width 73 (73—339) u; width of head-capsule 26 (21—47) u, between antennae 12
(11—14) u, n = 76; dorsal spinule-rows 4 (4—0) + 9 (7—10), ventrals 4 (4—0) + 8
(7—9), n = 10; respiration metapneustic. Second instar (fig. 70); length 664
(537—1296), width 269 (269—664) u; width of head-capsule 65 (45—68), between
antennae 22 (16—22) u, n = 50; dorsal spinule-rows in posterior part absent, in
anterior 4 (4—0), ventrals in posterior part absent either, in anterior part 8 (8—0),
n = 10. Third instar (figs. 71—73); length 1422 (901—1801), width 948 (584— 1138)
u; width of head-capsule 71 (45—85), between antennae 23 (17—31) u, n = 63;
spathula sternalis bilobed (figs. 74, 75), width 24 (19—28), height 60 (52—64),
distance between points of lobes 14 (13—22), height of lobes 9 (8—9), height of
median incision 9 (S—9) u, n = 10. Pupa length 1228 (1212—1643), prothoracic
horns 126 (122—146), apical papillae 28 (28—43) u, n = 8; sensory-pores between
the corresponding papillae of the two pairs of dorsal rows (fig. 95).

Papillary pattern (fig. 24b). Dorsal collar papillae absent, one pair of ventral
collar papillae. On thoracic segments one pair of dorsal papillae, 2 pairs of pleural
papillae and (one or) two pairs of laterals; on Th 1 one pair of sternal papillae, on
Th 2 and 3 one pair of ventrals. On segments A 1—7 two pairs of dorsal papillae,
one pair of pleurals and one pair of laterals, one (or two) pairs (posterior ventral
papillae only distinct in first instar) of ventral papillae; on A 8 one pair of dorsal,
pleural and ventral papillae, lateral papillae absent; one pair of terminal papillae
on AS.

Gall (fig. 97) length 1.6—2.1 mm, n 1.8 mm, width 1.0—1.8 mm, n 1.3 mm,
n = 20; galling of the fruit; gall ovoid, more or less hairy, wings present, although
not so large as in healthy fruits; window-pit distinct (B. pubescens).

Type material. — The type of Cecidomyia betulae Winnertz was destroyed in
1945 during the bombing of the Zoological Museum of Bonn (Mohn and Ulrich, in
litt.), and no other specimens of the original series exist. In regard to the (wrong)
low number of antennal segments in the original description, there is a good
chance that the type-specimen Winnertz described, belonged to this species. The
other species that could have been in Winnertz’ material, all have more antennal
segments. Moreover, in Poland, where the type-locality is situated, this species
seems to be most common. A new type is selected: Neotype &', slide no. 760223.8,
The Netherlands, Renkum, 15.1.1975, ex fruit galls of B. pendula, Allotype 9, slide
no. 760206.3, The Netherlands, Meyendel, 28.iv.1975, ex fruit galls of B. pubescens,
leg. J. C. Roskam. Although this material does not originate from Poland, it comes
as near to the original type locality as in my case is practicable (cf. Code, article 75
c5).

The original host-plant, B. alba, is a polytypic species, comprising among others
B. pendula and B. pubescens. Paratypical material is presented to the Bonn
Museum.

J.C. Roskam: The genus Semudobia 169

Material (all my localities are from The Netherlands) in coll. Rijksmuseum van
Natuurlijke Historie, Leiden. Adults slides no. 760223.7—8, 760206.2—3,
760216.1, Renkum, 15.1.1975, ex fruit galls of B. pendula (3); Meyendel, 28.iv.1975,
ex fruit galls of B. pubescens (2). First instar larva slide no. 750519.1, Duivenvoor-
de, 19.v.1975, ex B. pubescens; second instar slide no. 750619.4, Duivenvoorde,
19.vi.1975, ex fruit galls of B. pubescens; third instar slide no. 751001.1, Meyendel
1.x.1975, ex fruit galls of B. pubescens; pupa slides no. 730402.1—2, 730410.1—4,
Meyendel, 2 and 10.iv.1973, ex fruit galls of B. pendula, leg. J. C. Roskam.

Semudobia brevipalpis sp. nov.
(male: figs. 16, 30, 31, 44, 45; female: figs. 38, 39, 51, 56, 57, 103; imm. stages: figs. 77—81, 98)

Male. — Width of head 1.32 (1.25—1.32) times its height. Antenna (figs. 30, 31)
with 2 + 13+ (12+—13+) segments, its length 2.45 (2.41—2.60) times the width of
the head; scape ventrally with 1 (O—1) very short, 5 (4—5) lateral and 1 (0—1)
latero-distal long setae; third flagellomere with its node 1.10 (1.04—1.15) times as
long as its diameter; internodes oblong, internode of third flagellomere 0.42
(0.42—0.48) times as long as the node. Maxillary palp 0.19 (0.19—0.22) times the
width of head; third segment 2.00 (1.73—2.20) times as long as the second; first
and second segments partially grown together. Fronto-clypeal setae 2 (2—5), 12
(S—13) setae on mesanepisternum, 11 (8—11) on mesanepimeron. Wing length
1.78 (1.52—1.78) mm, 2.0 (2.0—2.1) times as long as wide; RS 2.2 (2.1—2.2) times
as long as R1; Sc indicated by 4 (4S) sensory-pores, 4 (4—6) pores on basal part
and 3 (2—3) on distal part of RI, 3 sensory-pores on medio-distal portion of RS.
Tibia of hind leg 0.76 (0.72—0.87) times as long femur, second tarsal segment 0.44
(0.43—0.50) times this length; fifth tarsal segment 0.85 (0.80—0.91) times as long as
fourth. Distimere acute (fig. 44), 0.55 (0.45—0.56) times as long as basimere;
inferior lamella about half way U-shapedly incised, sides converging to apex, lobes
distally acute (fig. 45); aedeagus a little longer than the weakly chitinized, broadly
rounded triangular claspettes.

Female. — Width of head 1.32 (1.31—1.34) times its height. Antenna (figs. 38,
39) with 2 + 11+ (10—12) segments, its length 1.61 (1.56—1.67) times the width of
head; scape ventrally with 1 (O—1) very short, 4 (3—4) lateral and 1 (0—1) latero-
distal long setae; third flagellomere with its node 1.09 (1.09—1.25) times as long as
its diameter. Maxillary palp (fig. 103) 0.19 (0.19—0.24) times the width of head;
length of third segment 2.00 (1.55—2.22) times the length of second; first and
second segments partially grown together. Fronto-clypeal setae 2 (0—4), 4 (3—6)
setae on mesanepisternum, 6 (3—14) setae on mesanepimeron. Wing length 1.68
(1.68—1.84) mm, 2.18 (2.17—2.31) times as long as wide; R5 2.22 (2.10—2.22)
times as long as RI; Sc indicated by 4 (34) sensory-pores, 3 pores on basal and 2
(2—3) on distal part of RI, 3 on medio-distal portion of RS. Tibia of hind leg 0.83
(0.82—0.87) times as long as femur, second tarsal segment 0.46 (0.44—0.49) times
this length; fifth tarsal segment 0.89 (0.82—0.91) times as long as fourth. Tergum 8
triangular (fig. 51), the height of this sclerite a little more than its width. Ovipositor
(fig. 57) 0.71 (0.70—0.73) times as long as hind femur, for about the half retracted

170 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

into the eighth abdominal segment; vaginal furca not remarkably chitinized,
superior lamella about a third longer than high, inferior lamella a little shorter
than wide (fig. 56).

Immature stages. — Egg length n 314 u, n + 2 SD (95% of egg lengths)
between 297—331 u; width n 112 u. First, second instar larvae and pupae are not

collected. Third instar (figs. 77—79): length 1388 (1101—1404), width 1069 |

(718—1084) u, width of head-capsule 79 (61—83), between antennae 24 (20—28) u;
spathula sternalis very wide with two small lobes (figs. 80, 81); width 55 (42—71),
height 25 (16—28), distance between points of lobes 39 (28—52), height of lobes 3
(2—5), height of median incision 6 (S—8) u, n = 15. Pupae not collected.

Papillary pattern (fig. 24c) very similar to S. betulae. Lateral papillae present on
A 8, on AS 2—3 pairs of terminal papillae.

Gall (fig. 98) length 1.5—2.0 mm, n 1.8 mm, width 0.9—1.4 mm, n 1.2 mm, n =
20; galling of the fruit, the gall is obovate and glabrous, wings almost completely
reduced, window-pit distinct, usually developed on both ad- and abaxial side of the
gall (B. populifolia).

Type material. — Holotype &, slide no. 760220.4, Allotype 9, slide no.
760318.1. Canada, Quebec, natural forest along Trans-Canada Highway, 9.11.1975,
ex fruit galls of B. papyrifera, leg. W. F. Grant.

Material in coll. Rijksmuseum van Natuurlijke Historie, Leiden. Adults slides
no. 760220.1,3—4, 750209.10, 760213.1, 760218.2—3, same data as type material.
Third instar larva slide no. 761101.14, same data as type-material.

The species is quite close to S. betulae: the larvae and pupae have two pairs of
rows of dorsal papillae on the abdomen, also the adult antennae are quite similar.
Nevertheless there are striking differences, viz., in the larvae the broad spathula
with the very small lobes, in the adults the bristles in latero-distal position on the
scape, the very short second palp segment, the broadly emarginated inferior
lamella in the male and the short, plump superior lamella in the female genitalia.

Semudobia steenisi sp. nov.

(male: figs. 109, 110, 113, 114; female: figs. 106—108, 111, 112, 115; imm. stages: figs. 116—122)

Male. — Width of head 1.12 (1.09—1.16) times its height. Antenna (figs. 109,
110) with 2+14 segments, its length 3.39 (3.18—3.97) times the width of head;
scape ventrally with 4 (1—4) very short, 3 (3—4) lateral and no dorsal long setae;
third flagellomere with its node 1.00 (1.00—1.06) times as long as its diameter;
internodes oblong, internode of third flagellomere 0.63 (0.58—0.65) as long as the
node. Maxillary palp 0.24 (0.22—0.28) times the width of head, third segment 1.20
(1.12—1.35) times as long as second; first and second segments partially grown
together. Fronto-clypeal setae 2 (2—5); setae on mesanepisternum | (0—3), 2
(2—5) setae on mesanepimeron. Wing length 1.86 (1.64—1.91) mm, 2.29 (2.28—
2.46) times as long as wide; RS 2.11 (1.88—2.17) times as long as RI; Sc indicated
by 6 (4—7) sensory-pores, sensory-pores on basal part of RI indistinct and 2 on
distal part, 3 pores on medio-distal portion of R5. Tibia of hind leg 0.79
(0.76—0.91) times as long as femur and second tarsal segment 0.59 (0.50—0.64)
this length; fifth tarsal segment 0.62 (0.62—0.77) times as long as fourth. Distimere

J.C. ROsKAM: The genus Semudobia 171

(fig. 113) 0.46 (0.46—0.52) times as long as basimere; inferior lamella weakly
emarginate (fig. 114); aedeagus about as long as the chitinized, triangular, rather
slender claspettes.

Female. — Width of head 1.15 (1.12—1.15) times its height. Antenna (figs. 111,
112) with 2 + 12+ segments, its length 2.07 (2.03—2.33) times the width of head;
scape ventrally with 2 (2—4) very short, 3 lateral and no dorsal long setae; third
flagellomere with its node 1.06 (1.00—1.11) times as long as its diameter. Maxillary
palp (fig. 115) 0.31 (0.28—0.32) times the width of the head; length of third
segment 1.15 (1.06—1.20) times the length of the second; first and second
segments partially grown together. Fronto-clypeal setae 3 (2—3); 3 (2—5) setae on
mesanepisternum, 3 (2—4) setae on mesanepimeron. Wing length 1.86 (1.84—2.00)
mm, 2.48 (2.31—2.48) times as long as wide; R5 2.11 (2.11—2.28) times as long as
R1; Sc indicated by 4 (4—7) sensory-pores, 3—5 indistinct sensory-pores on basal
part of RI and 2 (2—3) on distal part, 3 pores on medio-distal portion of RS. Tibia
of hind leg 0.79 (0.77—0.82) times as long as femur, second tarsal segment 0.63
(0.61—0.70) times this length; fifth tarsal segment 0.77 (0.77—0.87) times as long as
fourth. Tergum 8 bluntly triangular to H-shaped (fig. 108), the height of this
sclerite about one-and-a-half times its width. Ovipositor (fig. 106) 0.77 (0.72—0.81)
times as long as hind femur, for one to about three-quarters retracted into the
eighth abdominal segment; vaginal furca not remarkably chitinized; superior
lamella 1.92 (1.71—1.92) times as long as high; inferior lamella a little shorter than
wide (fig. 107).

Immature stages. — Egg length and first instar larvae not studied. Second instar
(fig. 117) length 821, width 483 u; width of head-capsule 47, between antennae 17
u; dorsal spinule rows in posterior part absent, in anterior part 3; ventrals in
posterior part absent too, in anterior part 7; n = 1. Third instar (figs. 118—120)
length 1563 (1420—1723) u, width 1101 (909—1196) u; width of head-capsule 57
(52—86), between antennae 30 (28—39) u, spathula sternalis (figs. 121, 122)
bilobed, it may form one structure or consist of two independent pieces; width 31
(28—52), height 22 (19—42), distance between points of lobes 11 (11—39), height
of lobes 6 (5—8), height of median incision 8 (6—9) u, n = 16. Pupa length 1372
(1324—1627) u; prothoracic horns 79 (71—94) u; apical papillae 30 (25—36) u;
n=4.

Papillary pattern (fig. 24d) rather similar to S. betulae. Only one pair of lateral
papillae on thoracic segments and two pairs of ventrals on segments A 1—7; on AS
2—3 pairs of terminal papillae.

Gall (fig. 116) length 2.1—3.0 mm, n 2.6 mm, width 1.4—2.1 mm, n 1.8 mm, n =
20; galling of the fruit; the gall is obovate and usually concave on ad-axial side,
more or less hairy, wings present, although not so large as in healthy fruits;
window-pit distinct (B. occidentalis).

Type material. — Holotype &, slide no. 761101.4, Allotype ©, slide no.
761101.10. U.S.A., Wyoming, Fremont County, Wind River, 10 miles north of
Dubois, 17.1x.1976, ex fruit galls of B. occidentalis, leg. J.C. Roskam.

Material in coll. Rijksmuseum van Natuurlijke Historie, Leiden. Adults slides
no. 761101.1—3, 5—9, 11—12, same data as type-material. Second instar larva

172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

slide no. 730411.3, Canada, Br. Columbia, O’Kanagan Valley, 13.viii.1959, ex fruit
galls of B. occidentalis, leg. C. G. G. J. van Steenis; third instar slide no. 761101.13,
same data as type material; pupa slide no. 730413.13, U.S.A., Montana, Bridger
Canon, alt. 1600 m, 16.ix.1905, ex fruit galls of B. occidentalis, leg. Blankinship 460.

I have great pleasure in naming this species after prof. dr. C. G. G. J. van
Steenis, Rijksherbarium, Leiden. He collected the herbarium specimen of river
birch in which I found larvae of this species. Together with S. betulae and S.
brevipalpis, S. steenisi belongs to the “betulae species complex”. All these species
induce fruit galls with window-pits and have larvae with two pairs of rows of dorsal
papillae on the A 1—7 segments. S. steenisi is the only species of this complex with
only one pair of rows of lateral papillae on the thoracic segments and two pairs of
rows of ventrals on the abdominal ones. The pupae have short prothoracic horns.
Especially in the males the long antennae with very long internodes are
remarkable; in females the length of the superior lamella of the ovipositor is just
intermediate between S. betulae and S. brevipalpis.

Only at a very late moment during this study I recognised West-Nearctic midges
of fruit galls to be different from East-Nearctic ones. Before my collection trip to
the U.S.A. in 1976 I considered all the larvae derived from Nearctic fruit galls to
belong to one, very variable, species. After this trip I reared adults of West-
Nearctic material. These adults were different from East-Nearctic midges in many
characters. Moreover, it was possible to correlate these differences with
differences in larval characters. Because the plates of the other Semudobia species
were completed before I depicted S. steenisi, all the figures of this species are
united in the final two plates of this paper.

Semudobia tarda sp. nov.
(male: figs. 28, 29, 46, 47; female: figs. 36, 37, 52, 58, 59, 104; imm. stages: figs. 9, 62, 63, 88—94, 99)

Male. — Width of head 1.23 (1.18—1.26) times its height. Antenna (figs. 28, 29)
with 2+ 14 segments, its length 2.81 (2.65—2.81) times the width of head; scape
ventrally with 3 (2—3) very short, 4 (3—4) lateral and 0 (0—6) dorsal long setae;
third flagellomere with its node 1.11 (0.96—1.11) times as long as its diameter;
internodes subquadrate to oblong, internode of third flagellomere 0.48 (0.46—
0.56) as long as the node. Maxillary palp 0.28 (0.24—0.28) times the width of head,
third segment 0.85 (0.74—0.88) times as long as second. Fronto-clypeal setae 3
(2—3); setae on mesanepisternum absent, 3 (3—7) setae on mesanepimeron. Wing
length 1.9 (1.5—1.9) mm, 2.1 (2.1—2.2) times as long as wide; RS 2.4 (2.1—2.5)
times as long as R1; Sc indicated by 5 (S—7) sensory-pores, 4 (4—6) sensory-pores
on basal part of RI and 2 on distal part, 3 pores on medio-distal portion of RS.
Tibia of hind leg 0.96 (0.76—0.99) times as long as femur, and second tarsal
segment 0.53 (0.47—0.55) times this length; fifth tarsal segment 0.79 (0.74—0.88)
times as long as fourth. Distimere (fig. 46) rather inflated, 0.48 (0.48—0.49) times
as long as basimere; inferior lamella about half-way narrowly V-shapedly incised

J.C. ROSKAM: The genus Semudobia 173

(fig. 47); aedeagus rather slender, a little longer than the rectangular, distally
emarginated and often heavily chitinized claspettes.

Female. — Width of head 1.25 (1.23—1.30) times its height. Antenna (figs. 36,
37) with 2 + 12+ (11—13) segments, its length 1.76 (1.67—1.98) times the width of
head; scape ventrally with 2 (2—3) very short, 1 (1—4) lateral and 1 (O—3) dorsal
long setae; third flagellomere with its node about as long as (1.00 (1.00—1.08)
times) its diameter. Maxillary palp (fig. 104) 0.24 (0.24—0.29) times the width of
head; third segment 1.12 (1.00—1.27) times as long as second. Fronto-clypeal setae
2 (O—4); setae on mesanepisternum absent or indistinct, 6 (4—6) setae on
mesanepimeron. Wing length 2.1 (1.8—2.1) mm, 2.1 (2.1—2.2) times as long as
wide; RS 2.4 (2.3—2.4) times as long as R1; Sc indicated by 4 (4—6) sensory-pores;
4 (4—6) sensory-pores on basal part of Ri and 2 on distal part, 3 pores on medio-
distal portion of R5. Tibia of hind leg 0.94 (0.91—0.94) times as long as femur and
second tarsal segment 0.51 (0.47—0.52) times this length; fifth tarsal segment
about as long as fourth. Tergum 8 triangular (fig. 52), the height of this sclerite
about equal to its width. Ovipositor (fig. 59) 1.31 (1.09—1.31) times as long as hind
femur, for one-third to about half-way retracted into the eighth abdominal
segment; oviduct surrounded by a remarkably chitinized vaginal furca; superior
lamella 1.5 (1.5—1.6) times as long as high; inferior lamella very short and truncate
(fig. 58).

Immature stages. — Egg length n 255 u, n + 2 SD (95% of egg lengths)
between 235—275 u; width n 72 u. First instar larva (figs. 62, 63); length 255
(212—539), width 97 (85—303) u; width of head-capsule 29 (26—34), between
antennae 12 (11—14) u, n = 17; dorsal spinule-rows 2 (2—0) + 8 (6—8), ventrals |
(2—0) + 7 (6—7), n = 10; respiration apneustic. Second instar (fig. 88); length 632
(506— 1248), width 269 (237—679) u; width of head-capsule 65 (47—71), between
antennae 17 (17—22) u,n = 14; dorsal spinule-rows 0 (—1) + 5 (0—8), ventrals 0
(O—2) + 11 (9—11), n = 10. Third instar (fig. 89—91); length 1596 (948—1691),
width 1185 (711—1185) u; width of head-capsule 73 (59—90), between antennae 19
(19—28) u, n = 31; spathula sternalis large, bilobed (fig. 92, 93), width of spathula
71 (27—71), height 89 (66—102), distance between points of lobes 47 (20—47),
height of lobes 27 (19—27), height of median incision 17 (9—17) u; n = 10. Pupa
length 1627 (1404—1643), prothoracic horns 141 (133—143) and apical papillae 47
(38—53) u, n = 7; sensory-pores situated laterally of the corresponding papillae of
the dorsal row (fig. 94).

Papillary pattern (fig. 24a). Dorsal collar papillae absent, one pair of ventral
collar papillae. On thoracic segments one pair of dorsal, two pairs of pleural, and
one pair of lateral papillae; on Th 1 one pair of sternal papillae, on Th 2 and 3 one
pair of ventrals. On segment A 1—7 only one pair of dorsal papillae, one pair of
pleural, lateral and ventral papillae; dorsal papillae are absent on A 8, there is one
pair of pleural papillae, one pair of laterals and one pair of ventrals; one pair of
terminal papillae on AS.

Gall (fig. 99) length 1.7—2.6 mm, n 2.1 mm, width 1.7—2.4 mm, n 2.0 mm, n =
20; galling of the fruit; the fruit is button-like swollen, subrotund, wings are
completely or nearly completely reduced; window-pit rather indistinct (B.
pubescens).

174 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

Type material. — Holotype 4, slide no. 760223.11, The Netherlands, Renkum,
15.1.1975, ex fruit galls of B. pendula. Allotype 9, slide no. 760211.2, The
Netherlands, Meyendel, 28.iv.1975, ex fruit galls of B. pubescens, leg. J. C.
Roskam.

Material (all my localities are from The Netherlands) in coll.Rijksmuseum van
Natuurlijke Historie, Leiden. Adults slides no. 760223. 9—10, 760211.1 and 11,
males with the same data as the holotype, females as allotype. First instar larva
slide no. 750519.3 Duivenvoorde, 19.v.1975, ex female catkins of B. pubescens;
second instar slide no. 750619.4, Meyendel, 22.vi.1972, ex fruit galls of B.
pubescens; third instar slide no. 751001.2, same locality as second instar, 1.x.1975;
pupa slides no. 730402. 4—6, 9, same locality as second instar, 2.1v.1973, leg. J. C.
Roskam. As regards the phenology (see below) this species is the latest, hence the
name tarda, the retarded. In the sense of Hennig (1966), this is the Semudobia
species with many apomorphic characters. According to Mohn (1955), three pairs
of rows of dorsal papillae seem to be most common in gall midge larvae, and one
pair of rows in S. tarda therefore must be considered apomorphic. This is the only
Semudobia species with apneustic first instar larvae. The heavily chitinized genital
structures, viz., claspettes in male and vaginal furca in female, only occur in this
species. In my opinion, these character stages must also be regarded as
apomorphic.

Semudobia skuhravae sp. nov.

(male: figs. 26, 27, 48, 49; female: figs. 34, 35, 53, 60, 61, 105; imm. stages: figs. 11—13, 66, 69, 82—87,
96, 100)

Male. — Width of head 1.08 (1.08—1.22) times its height. Antenna (figs. 26, 27)
with 2+ 13 (12—13) segments, its length 2.32 (2.26—2.49) times the width of head;
scape ventrally with one very short, 3 (2—4) lateral and 0 (0—2) dorsal long setae;
third flagellomere with its node 1.21 (1.08—1.26) times as long as its diameter;
internodes transverse to subquadrate, internode of third flagellomere 0.31
(0.31—0.41) times as long as the node. Maxillary palp 0.28 (0.23—0.31) times the
width of head, length of third segment 1.53 (1.05—1.57) times the length of second.
Fronto-clypeal setae 4 (1—5); 8 (0—8), often indistinct, setae on mesanepister-
num, 8 (4—9) on mesanepimeron. Wing length 2.14 (2.08—2.14) mm, 2.2 (2.2—2.3)
times as long as wide; RS 2.2 (2.2—2.4) times as long as RI; Sc indicated by 6
(6—9) sensory-pores, 7 (5—7) pores on basal part of Rl and 2 on distal part, 3
pores on medio-distal portion of RS. Tibia of hind leg 0.84 (0.79—0.87) times as
long as femur and second tarsal segment 0.52 (0.47—0.57) times this length; fifth
tarsal segment 0.73 (0.73—0.92) times as long as fourth. Distimere (fig. 48) 0.55
(0.52—0.56) times as long as basimere; inferior lamella for about one-third broadly
V-shaped incised (fig. 49); aedeagus usually shorter than the verrucose triangular
claspettes.

Female. — Width of head 1.19 (1.09-—1.19) times its height. Antenna (figs. 34,
35) with 2 + 12 (11*—13) segments, its length 1.61 (1.61—1.87) times the width of
head; scape ventrally with 1 (1—2) very short, 3 (3—4) lateral and 4 (2—9) dorsal
setae; third flagellomere with its node 1.08 (1.08—1.18) as long as its diameter,

J.C. ROSKAM: The genus Semudobia 175

ultimate flagellomere rounded. Maxillary palp (fig. 105) 0.23 (0.23—0.28) times the
width of head; length of third segment 1.33 (1.18—1.69) times the length of second.
Frontoclypeal setae 5 (1—7); 6 (1—8), often indistinct setae on mesanepisternum,6
(5—9) setae on mesanepimeron. Wing length 2.14 (2.08—2.14) mm, 2.2 times as
long as wide; RS 2.3 times as long as RI; Sc indicated by 4 (4—7) sensory-pores,
about 6 pores on basal part and 2 (2—3) on distal part of RI, 3 sensory-pores on
medio-distal portion of RS. Tibia of hind leg 0.87 (0.87—0.91) times as long as
femur and second tarsal segment 0.54 (0.51—0.55) times this length; fifth tarsal
segment 0.79 (0.79—0.85) times as long as fourth. Tergum 8 rectangular to H-
shaped (fig. 53). Ovipositor (fig. 61) 0.76 (0.75—0.80) times as long as hind femur,
for about one-quarter to half-way retracted into the eighth abdominal segment;
vaginal furca not remarkably chitinized; superior lamella 2 times as long as high;
inferior lamella rounded, about as long as wide (fig. 60).

Immature stages. — Egg length n 359 u, n + 2 SD (95% of egg lengths) between
335—383 u, width n 115 u. First instar larva (figs. 66—69); length 333 (261—612),
width 103 (97—339) u; width of head-capsule 29 (26—37), between antennae 17
(12—17) u; dorsal spinule-rows 4 (3—4) + 8 (7—8), ventrals 4 (3—4) + 7 (7—8),n
= 10; respiration metapneustic. Second instar (fig. 82); length 679 (442—1059),
width 332 (221—727) u; width of head-capsule 67 (48— 79), between antennae 21
(16—26) u, n = 39; dorsal spinuie-rows absent, ventrals 0 (2—0) + 6 (6—8), n =
10. Third instar (fig. 83—85); length 1675 (1138—2022), width 932 (758—1329) u;
width of head-capsule 85 (67—112), between antennae 23 (23—34) u, n = 41;
spathula sternalis extremely variable (figs. 86, 87), more or less bilobed with a
variable number (0—6) of small teeth between-the usually larger, pointed, lobes,
width of spathula 17 (17—44), height 14 (14—20), distance between points of lobes
11 (11—35), height of lobes 3 (3—9), height of smaller, median teeth 2 (2—3) u, n
= 10. Pupa length 1547 (1117—1786), prothoracic horns 114 (114—160) and apical
papillae 42 (38—60) u, n = 10; sensory-pores between the corresponding papillae
of the second and third pairs of dorsal rows (fig. 96).

Papillary pattern (fig. 24e). One pair of dorsal collar papillae and two pairs of
ventral collar papillae on SS (only visible in first instar). On thoracic segments two
pairs of dorsal papillae, two pairs of pleurals, two pairs of laterals; on Th 1 one pair
of sternal papillae and on Th 2 and 3 one pair of ventrals. On segments A 1—7
three pairs of dorsal papillae, one pair of pleurals, one pair of laterals, two pairs of
ventral papillae; on A 8 one pair of dorsal, one pair of pleural, one pair of lateral,
and one pair of ventral papillae; 2—3 pairs of terminal papillae on AS.

Gall (fig. 100) length 1.4—2.0 mm, n 1.8 mm, width 1.0—1.2 mm, n 1.1 mm, n =
20; galling of the “bract”; the ovoid gall is situated between the spindle of the
catkin and the scale; window-pit absent (B. pubescens). In Japanese (Honshu,
Azegate, B. ermanii) and American (South Dakota, Black Hills, B. occidentalis)
material S. skuhravae is found in fruit galls. These galls have no window-pit and are
coalescent with the scale. When gall and scale are separated, the membraneous
cocoon of the larva becomes visible.

Type material. — Holotype &, slide no. 760223.4, Allotype 9, slide no.

760209.1, The Netherlands, Renkum, 15.1.1975, ex bract galls of B. pendula, leg. J.
C. Roskam.

176 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

Material (all my localities are from The Netherlands) in coll. Rijksmuseum van
Natuurlijke Historie, Leiden. Adults slides no. 760223.3, 5—6, 750115.2—3,
760209.2, 760216.2, same data as type material. First instar larva slide no. 750519.6,
Duivenvoorde, 19.v.1975, ex catkins of B. pubescens; second instar slide no.
750619.7, same locality as first instar, 19.vi.1975, ex bract galls of B. pubescens;
third instar slide no. 720713.2, Meyendel, 13.vii.1972, ex bract galls of B. pubescens.
Pupa slide no. 730328.2—3, same locality as third instar, 28.111.1973, leg. J. C.
Roskam.

This species is named after dr. Marcela Skuhrava, Prague, Czechoslovakia. She
was first (Skuhrava & Skuhravy, 1960, 1963) in suggesting the species to be
different from Semudobia betulae, remarking the correlation of this midge with the
bract gall. Moreover she was the first to depict the spathula. In this species all
three pairs of rows of dorsal papillae are developed on A 1—7, and so it is most
similar to the usual state of papillary pattern in Cecidomyiidae (Mohn, 1955).
Because all other Semudobia species have less than three pairs of rows, three pairs
of rows of dorsal papillae must be considered a plesiomorphic character. On the
ventral surface of larvae of S. skuhravae the papillary pattern is also very complete:
always two pairs of rows of lateral papillae on thoracic segments and two pairs of
rows of ventrals on the abdomen. In my opinion these characters of lateral and
ventral papillae must be considered a primitive condition in the genus. If we
combine the plesiomorphic characters with character states of other sema-
phoronts, we must conclude that relatively big eggs (this results in a relatively
small number of egg per female: about 50 in S. skuhravae, about 120 in S. tarda)
may also indicate a primitive condition. In this way, the galling of the bract may be
a primitive condition, too. In adults the situation is more problematic. Gagné
(1976) stated that a relatively large number of flagellomeres should be an
apomorphic character. In Semudobia relatively large numbers of flagellomeres
occur in S. skuhravae, with plesiomorphic characters in the larval papillary pattern,
and in S. steenisi, which shares apomorphic characters with S. betulae and S.
brevipalpis in the larval stages.

DISTRIBUTION

Semudobia midges depend entirely on Betula species. Therefore, their geograph-
ical distribution must be expected to be wholly or partially the same as that of
Betula. Betula occurs in the whole boreal and temperate part of the Holarctic
region (Meusel et al., 1965). For such a huge area, even after years of sampling,
the picture can only be very incomplete. Therefore, many more data are needed
to get a true understanding of the distribution of Semudobia species. Much
material for the present study was collected in western Europe. A collecting trip to
the U.S.A. resulted in many data on the distribution of Semudobia, especially in
the Black Hills and the Rocky Mountains. During a year, a graduate student had
the opportunity to study insects of birch catkins in Japan. Several samples of birch
catkins were collected by colleagues in Canada and Eastern Russia. Finally, many
of the present data are based on infections found in plant specimens from the
collections of the Rijksherbarium, Leiden and the Herbarium Vadense, Wage-
ningen.

J.C. ROSKAM: The genus Semudobia

177

S. skuhravae S. tarda 5. betulae S. brevipalpis S. steenisi

America East-Canada Montreal = - - + 4
Ontario - = + de da

Quebec - + = 4 Hy

West-Canada Alberta + - - = 2

Br. Columbia = - - = de

East-U.S.A. Connecticut + - - + =
Pennsylvania - - + + =

Washington DC > = - + =

West-U.S.A. Colorado + - - - +

Montana + + - = +

South Dakota + - - = 5

Wyoming + - - = +

Europe Austria + + = È
Belgium + = =
Czechoslovakia + + + = =

Denmark + + + - =

France + + + = =

Germany + + + - =

Great Britain + + + = &

Ireland + + + = =

The Netherlands + + = È

Norway + + + = È

Poland + + + - =
Soviet-Union Latvia + + + = =

Sweden + + + = =
Switzerland + + + = =

Asia Japan + + + = =

Table 1. Geographical distribution of Semudobia.

Soviet-Union Baikal Lake
Kamtschatica
Kirgistania
Nerchinsk

Wladiwostok

Table 1 shows a mainly Palaearctic distribution for S. betulae and S. tarda. When
I collected these gall midges in the U.S.A., this was always on introduced
European B. pendula. Also in Canada, where S. betulae and S. tarda were collected
on B. populifolia and B. papyrifera, respectively, an introduction of infected fruits of
Palaearctic birch species is supposed.

In North America two species of the “S. betulae complex’ are present: S.
brevipalpis in the eastern states of Canada and the U.S.A.; S. steenisi in the
western states including Rocky Mountains. A similar vicariance occurs in the host
plant. European Betula pubescens is replaced in eastern North America by B.
papyrifera and in western North America by B. occidentalis. The vicariance in

178

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

È number % infected

station S. betulae S. tarda S. skuhravae 5 betulae
of galls fruits S. tarda

a

Kootwijk 1 199 109272: 155 10 34 15550
2 WAS) 10930 614 101 = 6.08
3 495 DoS 418 40 37 10.45
4 2945 5.26 2397 248 300 9.67
5 369 558 253 112 4 2.26
6 126 0.72 110 3 13 36.67

Meyendel 1 486 Sea 293 125 68 2.34
2 545 5.50 226 159 160 1249
3 2323 15.43 2100 204 19 10.29

Duivenvoorde 2045 Saal 905 551 589 1.64

Binnen-Buiten

Liede 1174 6205 158 337 84 2023

Nieuwkoop 5742 11.82 3099 2259 384 109877

Ilperveld 1 1011 5.10 468 537 6 0.87
2 842 2.66 454 386 2 1.18

b

Kootwijk 1 138 0.37 110 24 4 4.58
2 558 ZIA 370 80 108 4.63
3 161 0.28 124 12 25 10233
4 367 OR Wal 264 98 5 2.69
5 1210 2.54 1035 164 11 6.31
6 190 0.70 172 6 12 28.67

Meyendel 1 1374 SO 833 443 98 1.88
2 1042 BROS 795 223 24 3:57
3 5752 10.87 3777 1709 266 DEA

Duivenvoorde 3274 4.70 873 2310 91 0.38

Binnen-Buiten

Liede 949 2.43 386 518 45 0.75

Nieuwkoop 5312 6.13 2329 2853 130 0.82

Ilperveld 1 3817 6.43 1770 2041 6 0.87
2 875 1.98 325 533 17 0.61

Table 2. Occurrence of Semudobia species in different study areas during the year

1976. a, early spring; b, late summer.

J.C. ROSKAM: The genus Semudobia 179

eastern Asia of B. pubescens by B. davurica seems not to have been followed by a
vicariance in Semudobia: all Japanese and East-Russian specimens of S. betulae (as
are those of S. tarda and S. skuhravae) are very similar to the European material.

It is of interest to note that S. betulae and S. tarda both are gall inducers in birch
fruits; they usually may be found together in the same birch populations. If
different habitats are sampled, the two species seem to be ecologically different. In
1976, the frequency of occurrence of Semudobia species in fourteen birch
populations of six areas was determined (Table 2). In each population about eight
trees were marked; in each sampling series a constant number of catkins were
collected. Two series were made, where possible from the same trees, one in early
spring just before the emergence of the midges and one in late summer, when
infections caused by this generation of midges could be easily recognised.

The study areas. — Kootwijk (52.11 N 5.46 E) is a very dry sandy area where B.
pendula is abundant. Meyendel (52.08 N 4.20 E) is a dune area with a rather
heterogeneous structure: B. pendula can usually be found on relatively dry sand
dunes, whereas B. pubescens is common in the wetter valleys. Duivenvoorde (52.06
N 4.24 E) is a rather wet area, overgrown with blackberry brakes and rushes; of
the two birch species, B. pubescens dominates here. Binnen-Buiten Liede (52.23 N
4.41 E), Nieuwkoop (52.10 N 4.50 E), and Ilperveld (52.29 N 4.58 E) are very wet
bogs; here again, B. pubescens is the dominant birch species. In the dry habitat
Kootwijk populations, 78% or more of the fruit infections is caused by S. betulae
(except Kootwijk 5, a young population). In wetter habitats (Duivenvoorde, Liede,
Nieuwkoop and Ilperveld), the S. tarda part of the fruit infections, especially in the
“b” series, is relatively large (44% or more). In the “scattered” Meyendel area
both infection “types” are represented with S. betulae predominant; there is no
distinct correlation with a wet or a dry habitat.

In Switzerland five birch populations at different altitudes were studied. In the
areas Kerenzerberg (47.07 N 9.08 E, alt. 600 m), Walenstadtberg (47.08 N 9.18 E,
alt. 800 m) and Boggenberg (47.06 N 9.01 E, alt. 1250 m) S. betulae, S. tarda and S.
skuhravae were all common; in the area Ahornen (47.05 N 9.00 E, alt. 1400 m), S.
tarda was absent and at Stausee Garichte (46.57 N 9.06 E, alt. 1600 m), S. betulae
was the only gall-inducing midge.

Latitude offers a similar pattern. In Norway, galls were collected in nine areas,
varying in North latitude from 58.33 to 64.30. In the two most northern localities,
Tunnsjö (64.45 N 13.00 E, alt. 750 m) and Kongsvoll (63.33 N 9.24 E, alt. 1000 m),
only S. betulae was present. In most other localities, both S. skuhravae and S.
betulae were common. The localities of this series where S. tarda was also present
are Aseral (58.33 N 7.24 E, alt. 250 m) and Ossjoën (61.20 N 11.30 E, alt. 500 m).

The reason why S. tarda is absent at high altitudes and/or latitudes may be a
phenological one. S. tarda is the latest in pupation (pag. 181); for this species the
vegetation period in these extreme areas may be too short to complete larval
development.

The southern-most locality in western Europe, where Semudobia infections
were found, is Saintes in France (45.45 N 0.45 W); S. betulae and S. tarda both are
abundant there. In Japan the most southern locality is Honshu, Azegate (36.44 N
139.27 E, alt. 1280 m); there S. skuhravae is the only gall inducer.

180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

In Europe, S. betulae, S. tarda and S. skuhravae regularly occur together; this
seems to be different in the Nearctic Region. In Canada and the U.S.A. 20
infected birch populations were sampled (populations with introduced Palaearctic
species excluded). In only 6 populations S. skuhravae was found together with
either S. brevipalpis or S. steenisi; in 7 populations S. skuhravae was the only gall
inducer, in 4 populations this was S. brevipalpis and in 3 populations S. steenisi. It is
of some interest to mention that in one large area (Black Hills, South Dakota) four
birch populations (B. occidentalis and B. papyrifera) could be detected with
Semudobia infections. In all these cases S. skuhravae was the only gall inducer.

LIST OF HOST PLANTS

Besides the fresh material, about 300 herbarium specimens with fruit catkins
were investigated on Semudobia infections. Kuzneva (in Komarov, 1936), Rehder
(1940), and Ohwi (1965) were used for the identification of birch taxa. The
nomenclature is according to Kruessmann (1960) and Fontaine (1970). The taxa
are listed in Table 3. Absence of infections by certain gall midge species of taxa
marked by an asterisk may be due to the small sample size.

The genus Betula may be divided into four sections. The section Acuminatae is
exclusively East Asiatic. The section Costatae has a disjunct areal: some of the
species are East Asiatic, other species are only found in the eastern part of North
America. The section Excelsae is Holarctic and the section Humiles is mainly
Circumpolar. None of the species belonging to the section Acuminatae were
infected at all. In the section Costatae infections are mainly common in the East
Asiatic species: B. costata, ermanii, grossa, jacquemontii and raddeana; here, S.
skuhravae is the most common gall inducer. In East American B. alleghaniensis
infections occur, but they are very rare (S. skuhravae, one gall in one cultivated
specimen only). The absence of infections in B. lenta may be explained by the very
typical odour of the cherry birch, which may act as a repellent compound for
Semudobia. Also in the section Humiles, S. skuhravae seems to be the predominant
species. B. nana deserves special attention. Bachmaier (1965) reported that
infections of Semudobia are not rare. In more than 30 specimens, however, I found
only one single gall of Semudobia in a Norwegian sample. In Einsiedeln,
Switzerland (47.07 N 8.45 E, alt. 900 m), a heather with B. nana is mixed with and
surrounded by normally infested B. pubescens; although flowering times of B. nana
and B. pubescens coincide (Hegi, 1957), no infections of Semudobia were found in
those dwarf birches. In Aseral, Norway (58.33 N 7.24 E), a bog with B. nana mixed
with B. pubescens and their hybrid B. x intermedia was visited. Both B. pubescens
and B. x intermedia were crammed with Semudobia infections. Here, again, not a
single gall was found in B. nana. Finally, infections are common in most species of
the section Excelsae.

PHENOLOGY

In The Netherlands (Meyendel), the respective ontogenetic stages were
observed in the following periods of 1976 (Table 4). In fig. 25, some differences in

|

J.C. RosKAM: The genus Semudobia 181

opapaynys *S
DPAIDI *S
anqnaag *S
181U0098 *S

Section Species

sıdjpdaaaaq *S

alnoides Buch.-Ham. - = = = =
cylindrostachya Lindl. apud Wallich - = - = 2
maxtmowteztana Regel

t+
be by bs

Acuminatae
(Regel).

>

albo-stnensts Burkill
alleghantensts Brit.
corylifolia Regel & Maxim.
costata Trautv.

ermantt Cham.

forrestit (W.W.Sm.) Hand.-Mazz.
grossa Sieb. & Zucc.
jacquemonttt Spach.

lenta L.

medwedtewtt Regel

nigra L.

raddeana Trautv.
schmidtii Regel

utilis D. Don

Costatae
(Regel).

+

I arl sp sr @ @- fl
La la +
I
Ù

I
+
I
I
i}

x
SII CI SLED So DI ES EEE SU Gs)

ar @ 0
1
I
1
I

chichibuensis Hara

x coerulea Blanchard
coerulea-grandis Blanchard
celtiberica Rothm. & Vasc.
davurica Pall.

x intermeara (Hartm.) Thomas
kirghisorum Sav.

litwinowti Doluch.
occidentalis Hook.
papyrifera Marsh.

pendula Roth

platyphylla Suk.

populifolia Marsh.

pubescens Ehrh.

ssp. carpathica, ssp. murithii
turkestanica Litv.

Excelsae
(Koch).

txt
SEIEN SE
Laat
1
I
\

+
SÈ

EE 95) 5) SD SS
+++0++++0++1
I++1++0 I
1++0++0104+4+ 1

I
I

Se

Humi les
(Koch).

apotensis Nakai

x borggraveana Zabel
fruticosa Pall.
glandulifera (Regel) Butler
glandulosa Michx.

humilis Schrank
middendorffii Trautv.

nana L.

pumila L.

. x purpusti Schneid.

. tatewakiana M. Ohki & S. Watanabe

La +e * OF
Ca) RG) Su EV I SI SLET SE

m

N spare U @ ar 0 sp @ U
I 1

te

Table 3. List of host plants. + infections under natural conditions; c, infections in
plants, cultivated outside their distribution area, e.g. botanical gardens; (*) less
than 5 specimens studied.

phenology of the Palaearctic species are demonstrated. S. skuhravae is the earliest
species to pupate, S. tarda the latest; S. betulae is the first in passing the first and
second stages.

Effects of different altitudes on phenology were studied in Switzerland. In
Walenstadtberg (alt. 800 m), situated on a sunny slope above the Walensee, no
remarkable differences with The Netherlands (Meyendel), were found. On

182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

Table 4. Phenology of Palaearctic Semudobia species.

stage period

third instar larva until 12.iv
pupa IS — Lv
adult 20.iv — 9.v
egg 28.iv — 15.v
first instar larva 8.v — 23.vi
second instar larva 9. vi — 12.vii

third instar larva from 16.vi

Boggenberg (alt. 1250 m), Semudobia appears about three weeks later, but in
Ahornen (alt. 1400 m) and at Stausee Garichte (alt. 1600 m) no eggs were observed
before the end of May, and at the end of July only first and second instar larvae
were found. Third instar larvae were collected in the last week of August at
Ahornen and Stausee Garichte.

Similar effects on the phenology are observed at high latitude. In the first half of
July, 1976, infections of Semudobia were observed in eight areas in Norway (61.20
N — 64.30 N). Most localities are situated at about 950 m alt. In six areas only first
instar larvae were present at that date; in only two cases, Ossjoén (61.20 N 11.30 E,
alt. 500 m) and Rindal (63.05 N 8.30 E, sea level), also second instar larvae could
be collected.

The full-grown larva of Semudobia undergoes diapausis; it is impossible to rear
adults unless they have passed a cold period. In one case (Wyoming, Wind River),
larvae of S. steenisi, collected in September, 1976, and, transferred to my room,
pupated and gave rise to adults in the beginning of October. A similar absence of
diapausal condition must have occurred in the material of Forsius (1927), who got
adults in August.

4
Eee
/
40 / 7
* ®
Va
20 A —— 0 S. skuhravae
NES # S. betulae
/
o @ ——e S. tarda
= —— | — 5
22 30 6 12 20
mn m Vv Vv WV

Fig. 25. Phenology of Semudobia. a, % pupated midges; b, % animals that passed first stage moulting; c,
% animals that passed second stage moulting.

J.C. ROSKAM: The genus Semudobia 183

ACKNOWLEDGEMENTS

It is with pleasure that I thank the following persons for their help in various
matters. At first my colleagues of the Division of Systematics and Evolutionary
Biology, especially prof. dr. R. Hegnauer, drs. M. Zandee, mrs. Lucy Brand, Stans
Kofman, mr. B. L. Wallaart; the graduate students mrs. Lydia Sevenster and miss
Gerda van Uffelen. Furthermore dr. R. J. Gagné, Washington; prof dr. P. G.
Gorovoi, Wladiwostok; prof. dr. W. F. Grant, Montreal; dr. K. M. Harris,
London; dr. G. H. Ives, Edmonton; dr. R. Lichtenberg, Vienna; prof. dr. E. Mohn,
Stuttgart; mr. W. Nijveldt, Wageningen; dr. O. Peck, Ottawa; dr. M. Skuhrava,
Prague; dr. H. Ulrich, Bonn. I am also grateful to the Netherlands Organization
for the Advancement of Pure Research (Z.W.O.), the Maria van Tusschenbroek
Fonds and the Uyttenboogaart-Eliasen Stichting, which made possible the part of
this study, done in Switzerland, Japan and the U.S.A.

REFERENCES

(page numbers between brackets refer to Semudobia and/or Betula)

Bachmaier, F., 1965. Die Insekten- und Milbenfauna der Zwergbirke. — Veröff. zool. St. Samml.,
Münch. 9: 55—158 (p. 82, 141).

Barnes, H. F., 1951. Gall Midges of Economic Importance. Part 5, Trees. 270 pp. (p. 111—114).

Bergenstamm, J. E. von & P. Low, 1876. Synopsis Cecidomyidarum. — Verh. zool.-bot. Ges. Wien 26:
1— 105 (p. 29—30).

Binnie, F. G., 1877. Further Notes on the Cecidomyidae, with descriptions of Three New Species. —
Proc. nat. Hist. Soc. Glasg. 3: 178—186.

Buhr, H., 1964. Bestimmungstabellen der Gallen (Zoo- und Phytocecidien) an Pflanzen Mittel- und
Nordeuropas. 1572 pp. (p. 251).

Collin, J. E., 1904. A list of the British Cecidomyidae arranged according the views of recent authors.
— Ent. mon. Mag. 40: 93—99.

Connold, E. T., 1901. British Vegetable Galls, an introduction to their study. 312 pp. (p. 30, 274, 295, pl.

120).

, 1909. Plant Galls of Great Britain. 292 pp. (p. 22, 80).

Docters van Leeuwen, W. M., 1957. Gallenboek. 2nd. ed. 332 pp. (p. 118—119).

Dobner, 1858. Lehrbuch der Botanik für Forstmanner. 2nd ed.: 300 [not seen].

Escherich, K., 1942. Die Forstinsekten Mitteleuropas. Ein Lehr- und Handbuch. Part 5 Hymenoptera
(Hautflügler) und Diptera (Zweiflügler). 746 pp. (p. 546—547, fig. 547).

Felt, E. P., 1908. Studies in Cecidomyiidae, II. — Bull. N. Y. St. Mus. 124: 307—422 (p. 357, 368).

, 1915. A study of Gall Midges, III. — Bull. N.Y. St. Mus. 180: 127—288 (p. 226—228).

, 1940. Plant Galls and Gall Makers. 364 pp. (p. 77).

Fontaine, F. J., 1970. Het geslacht Betula. — Belmontiana 13: 99—181.

Forsius, R., 1927. Cecidiologische Beiträge. — Mém. Soc. Fauna Flora Fenn. 1: 34—38.

Gabler, H., 1958. Kenntnis der Schadinsekten der Blüten und Samen von Forstgehölzen. — Arch.
Forstwes. 7: 786—827 (p. 793— 795).

Gagné, R. J., 1967. The genus Oligotrophus Latreille (Diptera: Cecidomyiidae) in North America and a
new species injurious to Betula papyrifera Marsh. — Ent. News 78: 129—134.

—, 1968. Revision of the Genus Asteromyia. — Misc. Publ. ent. Soc. Am. 6 (1): 1—40.

——., 1973. A Generic Synopsis of the Nearctic Cecidomyiidi (Diptera: Cecidomyiidae: Cecidomyii-
nae). — Ann. ent. Soc. Am. 66: 857—889.

—, 1976. New Nearctic records and taxonomic changes in the Cecidomyiidae (Diptera). — Ann.
ent. Soc. America 69 (1): 26—28.

184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

Harris, K. M., 1966. Gall midge genera of economic importance (Diptera: Cecidomyiidae). Part I: In- |
troduction and subfamily Cecidomyiinae; supertribe Cecidomyiidi. — Trans. R. ent. Soc.
Lond. 118: 313—358.

—— , 1968. A systematic revision and biological review of the cecidomyiid predators (Diptera: Ceci-
domyiidae) on world Coccoidea (Hemiptera: Homoptera). — Trans. R. ent. Soc. Lond. 119:
401—494,

Hegi, G., 1957. Illustrierte Flora von Mittel Europa. 2nd ed. Part 3 (1). 452 pp. (p. 141—163).

Hennig, W., 1966. Phylogenetic Systematics. 263 pp.

Hodges, S., 1969. Gall Midges (Diptera-Cecidomyiidae) and their Parasites (Hymenoptera) Living in
Female Birch Catkins. — Trans. Soc. Br. Ent. 18: 195-226.

Hollander, J. den, 1975. The growth of larvae of Tipula oleracea Linnaeus, 1758 (Diptera, Tipulidae). —
Tijdschr. voor Entomologie 118: 67—82.

Houard, C., 1908—1913. Les Zoocécides des Plantes d'Europe et du Bassin de la Mediterrannée. Part
1. 560 pp. (p. 192).

Inchbald, P., 1885. A year’s work among the gall-gnats. — The Entomologist 18: 36—39.

Judeich, J. F. & H. Nitsche, 1895. Lehrbuch der Mitteleuropaischen Forstinsektenkunde. Part 2. p.
737—1421 (p. 1390).

Kaltenbach, J. H., 1874. Die Pflanzenfeinde aus der Klasse der Insekten. 848 pp. (p. 609).

Kieffer, J. J., 1895. Beobachtungen über die Larven der Cecidomyinen. — Wien. ent. Ztg. 14: 1—17.

, 1898. Synopse des Cécidomyies d’Europe et d’Algerie decrites jusqu’a ce jour. — Bull. Soc.

Hist. nat. Metz (2)8: 1—64 (p. 22).

——, 1900. Monographie des Cécidomyies d'Europe et d’Algérie. — Ann. Soc. ent. Fr. 69:
181—472.

— _, 1913a. Glanures Diptérologiques. — Bull. Soc. Hist. nat. Metz (3)4: 45—55.

——, 1913b. Diptera. Fam. Cecidomyidae. — Genera Insectorum. Part 152: 52, pl. 3 fig. 19, pl. 13
fig. 11.

Kruessmann, G., 1960. Handbuch der Laubgehölze. Part 1. 495 pp. (227—239, pls. 61—62).

Kuster, E., 1911. Die Gallen der Pflanzen. 437 pp. (p. 78, 362, 365).

Kuzneva, O. I. in V. L. Komarov, 1936. Flora of the U.S.S.R. [English ed.] Part 5. pp. 593 (p. 213—241).

Liebel, R., 1889. Ueber Zoocecidien Lothringens. — Ent. Nachr. 15: 298-307.

Lintner, J. A., 1887. [Third] Report of the State Entomologist to the Regents of the University of the
State of New York for the Year 1886. — Rep. N.Y. St. Mus. 40: 83—197 (p. 85—86).

Low, F., 1878. Mittheilungen über Gallmücken. — Verh. zool.-bot. Ges. Wien 28: 387—406.

Macquart, 1853. Les arbres et arbrisseaux d'Europe et leurs Insectes. — Mém. Soc. Sc. Lille 1852,

_ Suppl. 1853: 25 [not seen].

Mamaev, B. M. & N. P. Krivosheina, 1965. Lichinki gallits (Diptera, Cecidomyiidae). Sravnitel’naya
morfologiya, biologiya, opredelitel’nye tablitsy. 276 pp. (p. 21).

Meusel, H., J. E. Jager & E. Weinert, 1965. Vergleichende Chorologie der zentraleuropäischen Flora.
Part 1, 583 pp. (p. 459—460). Part 2, 258 pp. (p. 118—119).

Mohn, E., 1955. Beitrage zur Systematik der Larven der Itonididae (=Cecidomyiidae, Diptera). Part |
Porricondylinae und Itonididae Mitteleuropas. — Zoologica, Stuttgart 105: 1—247 (p.
179—180, pl. 21, figs. 5—6).

, 1961. Gallmücken (Diptera, Itonididae) aus El Salvador. 4. Zur Phylogenie der Aspondyliidi
der neotropischen und holarktischen Region. — Senck. biol. 42: 131—330.

Ohwi, J., 1965. Flora of Japan [English ed.] 1067 pp. (p. 372—375).

Pitcher, R. S., 1957. The abrasion of the sternal spathula of the larva of Dasyneura tetensi (Rübs.) during

the post-feeding phase. — Proc. R. ent. Soc. Lond. (A)32: 83—89.

Porter, C. L., 1967. Taxonomy of Flowering Plants. 2nd. ed. 472 pp. (p. 224).

Rehder, A., 1940. Manual of cultivated trees and shrubs hardy in North America. 2nd ed. 996 pp. (p.
124134).

Ross, H., & H. Hedicke, 1927. Die Pflanzengallen (Cecidien) Mittel- und Nordeuropas, ihre Erreger
und Biologie und Bestimmungstabellen. 348 pp. (p. 103).

Rübsaamen, E. H., 1891. Mittheilungen über neue und bekannte Gallmücken und Gallen. — Z. Na-
turw. 64: 123—156 (p. 137—141).

J.C. RoskAM: The genus Semudobia 185

, 1892. Die Gallmücken des Kônigl. Museums für Naturkunde zu Berlin. — Berl. ent. Z. 37:
319—411.

Rübsaamen, E. H., & H. Hedicke, 1925—1939. Die Cecidomyiden (Gallmücken) und ihre Cecidien. —
Zoologica, Stuttgart, 29: 1—350 (p. 287, pl. 25, figs. 1—9).

Schiner, J. R., 1864. Fauna Austriaca. Part 2, Diptera: 381.

Skuhravä, M., 1970. Taxonomische Probleme der Gattung Clinodiplosis Kieffer 1894 und der Gattung
Trotteria Kieffer 1901 (Cecidomyiidae, Diptera). — Informationsbericht der Landwirtschaftli-
chen Hochschule Nitra- Biologische Grundlagen der Landwirtschaft no 8: 63—68.

Skuhrava, M. & V. Skuhravy, 1960. Bejlomorky. 271 pp. (p. 178—179, figs. 82 —83).

, 1963. Gallmücken und ihre Gallen auf Wildpflanzen. 116 pp. (p. 83—85, fig. 59).

——, 1964. Verbreitung der Gallmücken in Jugoslavien (Dipt., Itonididae). — Dtsch. ent. Zeit. N.S.
11: 449— 459,

Strassburger, E., F. Noll, H. Schenk & A. F. W. Schimper, 1971. — Lehrbuch der Botanik für Hoch-
schulen. 30st. ed. 842 pp. (fig. 688 L M).

Swanton, E. W., 1912. British Plant-Galls. A classified textbook of Cecidology. 285 pp. (p. 156).

Theobald, F. V., 1892. An account of British Flies. Part 1.215 pp. (p. 63).

Thomas, F., 1893. Die Mückengallen der Birkenfrüchte. — F.-n. Zeitschr. 2: 464— 465.

Tubeuf, C. von, 1893. Die Mückengallen der Birkenfrüchte. — F.-n. Zeitschr. 2: 463—464.

Wachtl, F. A., 1881. Entomologisch-biologische Studie. I. Beitrag zur Kenntnis der Lebensweise von
Cecidomyia betulae Wtz. — Mitt. forstl. Vers. Oesterreichs 2: 99— 101.

Walker, F., 1856. Insecta Britannica. Part 3. 372 pp. (p. 85).

Winnertz, J., 1853. Beitrag zur einer Monographie der Gallmücken. — Linn. Ent., Berl. 8: 154—322 (p.
234).

Yukawa, J., 1971. A Revision of the Japanese Gall Midges (Diptera: Cecidomyiidae). — Mem. Fac.

Agr. Kagoshima University 8(1): 1—203.

186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

Figs. 26—33. Fifth, sixth and ultimate antennal segments, 3. 26—27, S. skuhravae; 28—29, S. tarda;

30—31, S. brevipalpis; 32—33, S. betulae. c, circumfila; ss, sensorial spine.

J.C. RosKAM: The genus Semudobia 187

Figs. 34—41. Fifth, sixth and ultimate antennal segments, 9. 34—35, S. skuhravae; 36—37, S. tarda;
38—39, S. brevipalpis; 40—41, S. betulae.

188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

49

72

Figs. 42, 44, 46, 48, male genitalia, dorsal aspect; 43, 45, 47, 49, detail of superior and inferior lamella:

42-43, S. betulae, 44—45, S. brevipalpis, 46—47, S. tarda, 48—49, S. skuhravae. a, aedeagus; b.

basimere; c. claspette; d, distimere; il, inferior lamella; sl, superior lamella. 42, 44, 46, 48, x 220; 43, 45,
47,49, x 400.

J.C. Roskam: The genus Semudobia 189

Figs. 50—53, female eighth tergum; 54, 56, 58, 60, detail of female superior and inferior lamella, ventral

aspect; 55, 57, 59, 61, ovipositor, lateral aspect: 50, 54, 55, S. betulae; 51, 56, 57, S. brevipalpis; 52, 58,

59, S. tarda; 53, 60, 61, S. skuhravae. il, inferior lamella; sl, superior lamella; vf, vaginal furca. 50—53, x
120; 54, 56, 58, 60, x 260; 55, 57, 59, 61, x 85.

190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

Figs. 62—69. First instar larva. 62, 64, 66, head and thorax, dorsal aspect; 68, ditto, ventral aspect; 63,

65, 67, sixth to eighth abdominal segment and anal segment, dorsal aspect; 69, ditto, ventral aspect:

62—63 S. tarda; 64—65 S. betulae; 66—69 S. skuhravae; av, anterior ventral papilla; cd, dorsal collar

papilla; cv, ventral collar papilla; d, d1—3, dorsal papilla; ip, inner pleural papilla; 1, lateral papilla; op,

outer pleural papilla; p, pleural papilla; pv, posterior ventral papilla; s, sternal papilla; t, terminal
papilla; v, ventral papilla. x 400.

J.C. ROSKAM: The genus Semudobia 191

iS REN IE ID AO OSLO

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i
i
i

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75

Figs. 70—75. S. betulae. 70, head and thorax of second instar, ventral aspect; 71, head, supernumerary

segment and first thoracic segment of third instar, ventral aspect; 72, seventh and eighth abdominal

segments and anal segment of third instar, ventral aspect; 73, ditto, dorsal aspect; 74—75, spathula

sternalis: 74, The Netherlands, Meyendel, 75, Kamtschatca. For explanation of symbols, see figs.
62—69. 70—73, x 140; 74—75, x 425.

192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

x hd Tr ~ 4,

WR DEI ED > =
VERSO E 7
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TE = EXT
di d2 t SS x eee 23
<A

Figs. 77—81. S. brevipalpis. 77, head, supernumerary segment and first thoracic segment of third instar,

ventral aspect; 78, seventh and eighth abdominal segments and anal segment of third instar, ventral

aspect; 79, ditto, dorsal aspect; 80—81, spathula sternalis: 80, U.S.A., Washington DC., 81, Canada,
Quebec. For explanation of symbols, see figs. 62—69. 77—79, x 140; 80—81, x 425.

J.C. ROSKAM: The genus Semudobia 193

Figs. 82—87. S. skuhravae. 82, head and thorax of second instar, ventral aspect; 83, head,

supernumerary segment and first thoracic segment of third instar, ventral aspect; 84, seventh and

eighth abdominal segments and anal segment of third instar, ventral aspect; 85, ditto, dorsal aspect;

86—87, spathula sternalis: 86, The Netherlands, Meyendel, 87, Canada, Alberta. For explanation of
symbols, see figs. 62—69. 82—85, x 140; 86—87, x 425.

194 TIJDSCHRIFT VOOR ENTOMOLGGIE, DEEL 120, AFL. 6, 1977

N | Cf
SARI
d2 p 2 N i

ew Eten 5
SENS EEn Zat
SS << 7 SZ
NU
NZ

Figs. 88—93. S. tarda. 88, head and thorax of second instar, ventral aspect; 89, head, supernumerary

segment and first thoracic segment of third instar, ventral aspect; 90, seventh and eighth abdominal

segments and anal segment of third instar, vei:tral aspect; 91, ditto, dorsal aspect; 92—93, spathula

sternalis: 92, The Netherlands, Meyendel, 93, Japan, Sapporo. For explanation of symbols, see figs.
62—69. 88—91, x 140, 92—93, x 425.

J.C. Roskam: The genus Semudobia 195

di sp d2 ds

Figs. 94—96, seventh and eighth abdominal segments of pupa, dorsal aspect; 97— 100, gall, 101, healthy

fruit of Betula pendula; 102—105, female maxillary palp: 94, 99, 104, S. tarda; 95, 97, 102, S. betulae; 98,

103, S. brevipalpis; 96, 100, 105, S. skuhravae. dl—3, dorsal papilla; sp, sensory-pore. 9496, x 75;
97—101, x 13; 102—105, x 400.

196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 6, 1977

Figs. 106—116. S. steenisi. 106, ovipositor, lateral aspect; 107, detail of female superior and inferior

lamella, ventral aspect; 108, female eighth tergum; 109, male ultimate antennal segments; 110, male fifth

and sixth antennal segments; 111, female ultimate antennal segments; 112, female fifth and sixth

antennal segments; 113, male genitalia, dorsal aspect; 114, detail of male superior and inferior lamella;

115, female maxillary palp; 116, gall. 106, x 85; 107, 114—115, x 400; 108, x 120; 109—1 12, x 330; 113,
x 220; 116, x 13.

J.C. RoskaM: The genus Semudobia 197

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Figs. 117—122. S. steenisi. 117, head and thorax of second instar, ventral aspect; 118, head,

supernumerary segment and first thoracic segment of third instar, ventral aspect; 119, seventh and

eighth abdominal segments and anal segment of third instar, ventral aspect; 120, ditto, dorsal aspect;

121—122, spathula sternalis: 121, Canada, Br. Columbia, 122, U.S.A., Montana. For explanation of
symbols, see figs. 62—69. 117—120, x 140; 121—122, x 425.

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HARVARD

UNIVERSITY

TIJDSCHRIFT
VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

K. VEPSALAINEN and N. NIESER. — Life cycles and alary morphs of some Dutch
Gerris species (Heteroptera, Gerridae), p. 199— 212, figs. 1—2.

| Tijdschrift voor Entomologie, deel 120, afl. 7 Gupubliceerd 30-XII-1977 |

Scan

18
ak
oy, i A

LIFE CYCLES AND ALARY MORPHS OF SOME DUTCH
GERRIS SPECIES (HETEROPTERA, GERRIDAE)

by
KARI VEPSÄLÄINEN

Department of Genetics, University of Helsinki, Finland
and

NICO NIESER

Laboratorium voor Zoölogische Oecologie en Taxonomie, Utrecht, The Netherlands
With two figures

ABSTRACT

Material of Gerris species was collected in the Netherlands during one summer, mainly of G.
odontogaster (Zett.) and G. lacustris (L.), and less of G. argentatus Schumm. and G. thoracicus Schumm.
Dutch populations are bivoltine like those in Central Europe.

G. odontogaster and G. argentatus are long-winged in the overwintering generation and alary
dimorphic in the first new, midsummer generation. Non-diapause adults emerge during a period of one
month, in mid July at the latest. The last overwintered adults die about mid July, non-diapause adults
before the winter. Few eggs seem to be laid after mid August. The first new, overwintering diapause
adults (macropters) emerge about 10 July, and a few younger instar larvae are still observed in
September.

The life cycle of G. lacustris is similar to that of G. odontogaster, but short-winged adults also occur in
the overwintering generation. There is a notable peak of short-winged imagos in midsummer and
another peak in late summer. G. thoracicus is virtually long-winged throughout the year.

Evidence on colonization flights of midsummer non-diapause G. lacustris and G. thoracicus is given.

INTRODUCTION

Andersen (1973), Vepsäläinen (1971, 1974a, 1974b) and Vepsäläinen &
Krajewski (1974) have shown regularities in the relationship between the life
cycles and alary polymorphism of European Gerris species (see also Jarvinen,
1976).

1) Odontogaster type populations are univoltine and long-winged in northern-
most Europe, and multivoltine (mostly bivoltine) and dimorphic over the main
range of the species. Diapause is determined by shortening day lengths at the first
four larval instars. Hence adults which have moulted after mid July do not
reproduce before the following spring. In northern Europe immediately reproduc-
ing (non-diapause) adults are short-winged, while diapausing adults are long-
winged. In more southern populations the non-diapause group includes both short
and long-winged individuals. Overwintering adults are long-winged.

2) Lacustris type populations are univoltine in northernmost Europe, and

199

200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 7, 1977

bivoltine over the main range of the species. The populations are usually
polymorphic through the year, although morph frequencies fluctuate. Diapause
determination is as in odontogaster type populations. In northern bivoltine
populations immediately reproducing adults are mainly short-winged; in southern
populations both short and long-winged non-diapause individuals are found.

3) Rufoscutellatus and najas type populations are usually univoltine, and
monomorphically long-winged or wingless, respectively.

The above grouping is a simplification of the highly varying life cycles in
Finnish, Danish, Polish and Hungarian Gerris populations. For instance, the cycle
of G. thoracicus in southernmost Finland is of the odontogaster type but Hungarian
populations are, almost without exception, long-winged (Vepsalainen, 1974a,
1974b). Moreover, dimorphic G. najas De Geer populations are also known
(Poisson, 1957; Krajewski, 1969).

G. odontogaster and G. lacustris are distributed from northern Europe to the
Balkans, G. argentatus and G. thoracicus reach northern Africa (for a survey of
distributions, see Vepsäläinen, 1973). Diapause is thought to be an adaptation to
adverse conditions, i.e. in winter, at least in most parts of Europe. Indeed, even in
Faro, the extreme south of Portugal, the conditions in December are not
favourable for larvae to reach adulthood. This is due to low food supply and low
temperature. On the other hand, the autumn temperatures until November are
comparable to cool summer days in the Netherlands and Denmark. So for
southern populations a strategy with a longer reproductive season than in northern
Europe could be possible. This could apply even to Dutch populations compared
with Finnish, as in the Netherlands September is still quite warm, while in Finland
it is often too cold for Gerris development. As in the northern hemisphere the days
begin to shorten after summer solstice, about 22 June, diapause is determined all
over the species ranges at approximately the same time. Therefore the mechanism
of diapause determination may seem non-adaptive.

The purpose of the present work is to study the life cycles of Atlantic
populations to reveal possible differences in the diapause reaction between these
and northern and southern continental populations (Finland, Poland, Hungary).
Simultaneously we report the alary morph frequencies, as the conditions of ponds
and other small waters which are the main habitats of the Gerris species
investigated may, on the average, be more stable and predictable in Atlantic than
in continental climates. As a consequence Atlantic populations could maintain a
higher frequency of midsummer short-winged adults than continental populations.
The following are our results based on specimens collected in the Netherlands
during 1974.

SITES STUDIED

The study area (ca. 200 km?) is located in the northern part of the Veluwe, prov.
of Gelderland, with the village of Elspeet (52°17’N, 5°47’E) in its centre. The
landscape is dominated by some rows of sandy hills formed by pleistocenic ice,
one running approximately north—south from Nunspeet to Garderen. Most of our

VEPSÄLÄINEN & NIESER: Life cycles of Dutch Gerris species 201

sites are at the foot of these hills. Northwest of Heerde there is another chain of
such hills, the Woldberg.

Most of the area has been planted with coniferous trees, but there are still a
number of heathlands (e.g. the Speulderveld) and local stands of deciduous trees
(e.g. near Nunspeet) left.

Except for those on the Speulderveld, the sampled sites are quite isolated, lying
in small clearings in woods. With each locality description distances to nearest
sites sampled are given. All these aquatic habitats were isolated from each other
by land.

Our collecting sites are a subset of aquatic habitats of the Veluwe selected for
zooplankton and later Nepomorpha studies, and supplemented with a collection of
Speulderveld habitats. They were chosen in such a way that a maximal range of
local Gerris habitats could be sampled. The following localities were included:

NI, Nunspeet, Waskolk, about 1500 m?, surrounded by woods, isolated. Semi-aquatic vegetation (Ca-
rex, Eleocharis, Juncus effusus, Glyceria and Sparganium) at edges, from where most Gerridae were
collected. Bottom: sand with a thin layer of plant debris, depth about 1 m. Used as a swimming pool
during summer. Distance to N2, 4 km.

N2, Nunspeet, Ossenkolk, about 1000 m?, surrounded by woods, isolated. Vegetation at edges Eriopho-
rum, Juncus bulbosus and Sphagnum; pond filled with vegetation, mainly Eleocharis, Juncus bulbosus,
Sphagnum and Utricularia. Bottom: sand with a thin layer of plant debris, depth for the greater part
about 50 cm. Distance to NI, 4 km.

N3, Ermelo, Speulderveld, Carex rostrata pool, about 100 m?, in open heath, bordered by Molinia, with
Carex rostrata growing in the greater part of the pool. Bottom with a thick layer of plant debris,
depth of free water 10-50 cm. Distance to N7-15, 20-300 m.

N4, Garderen, Watergraafsmeertje, about 2500 m?, surrounded by woods, isolated. Juncus effusus and
Molinia at edges; pond for the greater part filled with vegetation, mainly Juncus bulbosus, Eleocharis
(multicaulis), Ranunculus and Scirpus. Bottom: sand with a thin layer of plant debris, depth for the
greater part about 20 cm. Distance to N3, 7-15, 6 km.

NS, Heerde, pond at Buitenzorg, about 500 m?, surrounded by woods, isolated. Edges: sand and dry
Sphagnum. For the greater part filled with vegetation, mainly Sphagnum. Bottom: sand with plant de-
bris, depth about 50 cm. Distance to N6, 3 km.

N6, Heerde, Kikkersgat, about 1000 m?, surrounded by woods, isolated. Edges with Juncus bulbosus,
Molinia, Rhynchospora, Scirpus and Sphagnum; pond for the greater part filled with Eleocharis multi-
caulis, Juncus bulbosus and Sphagnum. Bottom with a thick layer of plant debris, depth about 50 cm.
Distance to N5, 3 km.

N7-10, Ermelo, Speulderveld, the four pits, 2 to 10 m?, on open heath, separated from each other by
narrow dams, no water connections. Edges with Calluna and Molinia, pits partly filled with Juncus
bulbosus and overhanging withered Molinia. Bottom: loamy soil with some plant debris, depth 5-80
cm. Distance to N3. N11-15, 20-300 m. The population of each pit: NE = N7, SE = N8, NW = N9,
and SW = N10, has been studied separately.

N11, Ermelo, Speulderveld, Potamogeton pit, 40 m?, edges with Calluna, pit for the greater part filled
with Potamogeton natans. Bottom: loamy soil with plant debris, depth 10-100 cm. Distance to N3, N7-
15, 20-250 m.

N12, Ermelo, Speulderveld, Utricularia pit, 5 m?, on open heath, edges and bottom open sand with
plant debris, a few Utricularia minor in the pit. Depth 5-25 cm. Distance to N3, N7-15, 10-250 m.

N13, Ermelo, Speulderveld, Juncus articulatus pool, 20 m2, on open heath, edges with Calluna and Jun-
cus articulatus, pool for the greater part filled with Juncus bulbosus. Bottom: sand with plant debris,
depth about 20 cm but on 26.VI. it was without water, the bottom still wet; it is not known how many
days the habitat was dry. Distance to N3, N7-15, 30-200 m.

N14, Ermelo, Speulderveld, Salix pond, 60 m?, surrounded by Salix shrubs and some Typha; pond for
the greater part filled with Potamogeton natans and some Juncus bulbosus. Bottom: loamy soil with
plant debris, depth about I m. Distance to N3, N7-15, 15-300 m.

202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 7, 1977

N15, Ermelo, Speulderveld, Typha pond, about 200 m?, Gerridae collected in a corner overgrown with
Typha, about 20 m?. Bottom: sand with much plant debris (mainly from Typha), depth about 20 cm.
Distance to N3, N7-14, 15-300 m.

METHODS

The 15 populations were studied, and the larger populations partially collected,
every second week from 16 May to 4 September 1974, N1-6 already on 16-19
April. Predominantly adults were collected, but larvae of the fourth and fifth
instars were also taken to confirm reproduction. Occasionally younger larvae were
included. During sampling the sites were checked for the occurrence of first and
second instar larvae to estimate when the last eggs were laid. Unfortunately,
samples were not taken after 4 September when egg-laying perhaps still continued
on some of the sites.

Adults were classified according to wing length (short wings or wingless as
against long wings). Notes were made on age (on basis of hardening and
pigmentation grade of the chitin), colour of the female venter, stage of maturation
(ovarian and testis), presence of adipose tissue, pigmentation of the mesoalinotum,
and developmental stage of indirect flight muscles. When this information is
combined, it is possible to make a reliable study of the life cycles. Details and
reasons for the classification outlined here are given by Andersen (1973) and
Vepsalainen (1974a, 1974b).

Habitat classification

The sites mentioned above were grouped for habitat features according to
Vepsäläinen (1973) and showed little diversity (table 1). Shore vegetation was
either sparse or moderately dense, the water surface was totally or moderately
exposed to sunshine, and all ponds were stagnant and in late May over 10 cm
deep. Plant coverage on water surface was mostly moderately to very dense, and
showed in five cases a temporal change from scarce or moderate to dense. Two
times the succession was reversed.

One site was classified as permanent (risk of drying up negligible), seven as
semi-permanent (risk of drying up small), and seven as temporary (likely to dry up
during exceptionally prolonged hot periods, i.e. at least once in some 50 years). In
one semi-permanent and four temporary ponds depth of water had decreased
under 10 cm by 26 June. One site (N13) was dry on 26 June but again contained
water during the visit on 10 July.

RESULTS

Species numbers and habitats

Codes of each collecting site with habitat and species data are summarised in
table 1. The number of species (apart from G. rufoscutellatus) per site varies
between two and four (table 2). Permanent and semipermanent habitats (com-
bined) tend to harbour more species than temporary ones (P=.025, one-tailed

203

VEPSÄLÄINEN & NIESER: Life cycles of Dutch Gerris species

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204 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 7, 1977

Table 2. Number of species per habitat grouped for permanency.

Habitat number of species 2 3 4
Permanent |
Semi-permanent | | 5
Temporary 3 2 2

Mann-Whitney U-test; the null hypothesis is that the temporary habitat has the
same distribution over species numbers as the more permanent habitat group).

Notably G. argentatus seems to prefer (semi-)permanent habitats, being absent
only from one semi-permanent, but all temporary, habitats. G. lacustris seems to be
indifferent to this habitat feature. For the remaining species the data are
inconclusive.

Life cycles and alary morphs

We will try to give a general picture of wing length variation and life cycles in
Dutch Gerris. Hence we have summed the information from all 15 localities in figs.
l and 2. There are probably differences between populations (see Vepsäläinen,
1974b: 12-13) but our material is too scarce to reveal any.

4 15 + ls 37 38 }36 34 ye i

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80; \° |
60 {|

: |

20

30 12

APRIL MAY JUNE JULY AUGUST

a | 0 aM eo | MEM

Fig. 1. Seasonal changes in the proportions of n different female forms of Gerris odontogaster. The

numbers of females per sampling day are given above. The diagram is an approximation based on

dissected material of 84.4% of the micropter and 44.1% of the macropter females. The arrows above

indicate the first and last samples, respectively, where about newly emerged non-diapause females were

found. a) hard chitin, dark venter, eggs or maturing oocytes, macropter; b) as a but short-winged; c)

pale venter, eggs or maturing oocytes, macropter; d) as c but short-winged; e) dark venter, immature,
macropter; f) as e but short-winged.

1

n

VEPSÄLÄINEN & NIESER: Life cycles of Dutch Gerris species 205
Gerris odontogaster

The overwintering population is long-winged. The first fifth instar larva was
captured on 30 May. First short-winged adults were caught on 12 June and short-
winged individuals dominated from late June to early August. The specimens in
the late August and early September samples were almost all long-winged (fig. 1,
table 3).

The short-winged adults are classified as micropters, wing length usually varying
from | to 2 (see Vepsäläinen, 1974b, fig. 1). On 24 July, one male with wing length
3.5 (i.e. brachypterous) was collected (incompletely pigmented; age less than one
week). The midsummer micropters are non-diapausing individuals whose gonads
mature soon after the final moult. One to two weeks old micropterous females
usually have full-sized eggs. Some non-diapause macropters emerge simultan-
eously with micropters. Both non-diapause female types are usually easy to
distinguish from overwintered and diapause females by their pale venters in
otherwise completely pigmented individuals. Males, however, are almost without
exception dark below (for details, see Vepsäläinen, 1974a).

Diapause individuals with dark venter begin to emerge about 10 July, and
gradually this group replaces other groups. The last two non-diapause females,
which have been classified as semi-soft (refers to the hardening of the chitin), were
captured on 24 July. As the hardening of the chitin may take over two weeks after
the final moult, it can be estimated that the ecdysis has taken place about mid
July. After 24 July all soft or semi-soft females checked for maturing oocytes have
been in diapause. The mature, late summer females are older individuals, which
have already laid most egg batches and die before the winter.

On 4 September no young larvae (first to third instars) were observed on eight

Table 3. Fraction of short winged specimens for males and females in G. argentatus (apters), G. lacustris
(mainly brachypters) and G. odontogaster (micropters) with s (standard deviation in sample) and,
between brackets, n (the number of specimens in sample) over all habitats on the various sampling
days. Actually sampling took place on two consecutive days, of which the first is mentioned.

G. argentatus G. lacustris G. odontogaster

sexes pooled 3 © 3 Q
date
16.1V 0 (5) 0.07+0.07 (14) 0.10+0.10 (10) 0 (11) 0 ( 4)
16.V. 0 (18) 0.06+0.04 (32) 0.10+0.05 (42) 0 (12) 0 (15)
30.V. 0 ( 3) 0.11+0.06 (28) 0.32+0.09 (28) 0 (MiO ( 3)
IAAI 0 (5) 023208 (8) © (10) 0 (4) 0.38+0.17 ( 8)
26.VI. 0.75+0.22 ( 4) 0.47+0.13 (15) 0.67+0.10 (24) 0.96+0.04 (23) 0.97+0.03 (37)
10.VII. | (12) 042+0.08 (36) 0.41+0.08 (44) 0.73+0.10 (22) 0.95+0.04 (38)

24.VII. 0.80+0.18 ( 5) 0.27+0.06 (49) 0.29+0.07 (49) 0.74+0.10 (19) 0.61+0.08 (36)
7.VII. 0.39+0.14 (13) 0.12+0.04 (67) 0.15+0.05 (65) 0.64+0.13 (14) 0.65+0.08 (34)
21.VIII. 0 (11) 0.17+0.07 (30) 0.41+0.10 (27) 0.04+0.03 (47) 0.06+0.04 (47)
4.IX. 0 (29) 0.49+0.08 (39) 0.50+0.09 (32) 0.02+0.02 (65) 0.01+0.01 (114)

206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 7, 1977

sites, and in seven populations they were very few or few. The species could not be
identified in the field. On 21 August four populations had still “quite a number” of
young larvae.

The one macropterous male checked for testis maturation on 7 August still had
soft chitin (hence it was not over a week old) and well-developed testes. It is not
known which proportion of late summer males has well developed testes nor
whether they actually copulate in late summer.

Histolysing indirect flight muscles were already found on 16 April in over-
wintered females with developing oocytes, and totally histolysed ones in females
with eggs on 16 May. Micropters have undeveloped flight muscles, but non-
diapause as well as diapause macropters develop indirect flight muscles in
approximately two to three weeks in nature. It seems that an individual with the
combination of characteristics — fully pigmented, semi-hard, mesoalinotum III or
perhaps II (for the classes, see Andersen, 1973, fig. 10), and indirect flight muscle
fibres with a diameter of 35-50 um — is already capable of flight.

Adipose tissue was found in individuals with undeveloped gonads. Non-diapause
individuals did not have adipose tissue; this was also the case with the
micropterous female with eggs, collected on 4 September.

In the following we list some exceptions from the above life history picture. In
the 12 June samples we found one semi-soft/semi-hard (hence not overwintered),
fully pigmented, dark-ventered female with undeveloped oocytes, and one semi-
soft, almost fully coloured female with dark venter and undeveloped oocytes (both
macropterous). Usually the oocytes of new-generation females which develop in
early summer are already well maturing in fifth larval instars (e.g. Andersen, 1973,
fig. 7).

In the 10 July sample there was one semi-hard, completely pigmented,
macropterous female with light venter but totally undeveloped oocytes (meso-
alinotum II and diameter of indirect flight muscle fibres about 30 um).

Collected on 26 June, there is one light-ventered micropterous female with
abnormal intestine and no eggs, and on 10 July, one semi-soft, light-ventered
micropterous female with contents of abdomen “rotten”’.

One micropterous male, collected on 24 July, had asymmetrical wings, one
hemielytron being longer and deformed.

Sex ratio was female weighted (39.5% males, n=554, P<.001; .95 confidence
limits for fraction of males .36-.44). The deviation is a sum effect of both
midsummer micropters and late summer diapause macropters. (Heterogeneity x
= 13.60, .10<P<.05; with the null hypothesis of even sex ratio in the samples. The
heterogeneity of the whole material (early summer included) is statistically
significant: x2,,,= 24.32, P<.025). Significant heterogeneity indicates a shift in sex
ratio during the cycle.

Gerris argentatus

The number of specimens is small, 105 imagoes in all. The general picture is like
that in G. odontogaster (table 3). The overwintering population is long-winged, the

VEPSÄLÄINEN & NIESER: Life cycles of Dutch Gerris species 207

midsummer one dimorphic. The short-winged individuals are mostly apterous but
sometimes micropterous. On 10 July one newly emerged male with hemielytra 6 +
and hind wings 4+ (brachypterous) was taken.

In this species also a few long-winged, non-diapause adults occur together with
apters: one macropterous female on 7 August had completely hardened chitin,
light venter and full-sized eggs.

Midsummer non-diapause females differ from diapause females by their more or
less light venter. However, Dutch G. argentatus females are usually notably darker
than the equivalent type in G. odontogaster (the former as the pattern argentatus D’
and the latter as B’ or C’ odontogaster in fig. 3, Vepsalainen, 1974a). No differences
have been noticed in the ventral coloration between non-diapause short- and long-
winged females.

The last overwintered adult (with strongly histolysed indirect flight muscles) was
collected on 26 June. The first new generation apters were taken on the same day,
and the first fifth instar larva on 12 June. No soft non-diapause individuals were
collected after 10 July, and the first diapause macropters were not taken before 7
August. (On 24 July only five adults were captured.)

Indirect flight muscle histolysis was noted from 16 April onwards in over-
wintered individuals. In late summer, diapause imagoes had well-developed indi-
rect flight muscles and voluminous adipose tissue.

The fraction of males was 50% (n=105).

Gerris thoracicus

In this species no definite evidence of alary dimorphism was obtained. The
collected 61 imagoes were macropters with the exception of one male brachypter
in the 10 July sample.

The Dutch population, like in the two previous species, is bivoltine (at least
partially). The first fifth instar larva was captured on 12 June, and the first new,
non-diapause imagoes (14 39) on 26 June. The last overwintered specimen (13)
was caught likewise on 26 June. The last young non-diapause imago was obtained
on 10 July, and the first records of diapause individuals are from 24 July (two semi-
hard young individuals, one of which had reached stage IV mesoalinotum
coloration). The last female with eggs occurred on 21 August.

G. thoracicus differs from the previous two species in that both diapause and
non-diapause individuals are nearly entirely dark below. The ventral coloration is
therefore not indicative of a reproductive stage.

Another difference with the preceding species is evident: usually, histolysis of
indirect flight muscles does not take place in overwintered G. thoracicus. In the last
overwintered specimen (1 & on 26 June) indirect flight muscles were still full-
sized.

The one brachypterous male (semi-hard, totally pigmented, hemielytra 7.2, hind
wings 5.2; called submacropter by many previous investigators) had well
developed testes but undeveloped flight muscles.

The fraction of males was 46% (n=61).

208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 7, 1977

Gerris lacustris

This species has a more complicated life cycle as even the overwintering
population is polymorphic. A notable peak of brachypters (the other morph in this
species, medially short wings) occurs in midsummer. Later the relative frequency
of brachypters declines to rise again in the early September sample. In late spring
brachypter frequencies are at the lowest (fig. 2, table 3).

The high frequencies of brachypters in mid summer are explained by the fact
that a higher proportion ofthe non-diapause imagoes are short-winged (fig. 2). The
situation is analogous with G. odontogaster and G. argentatus which have a mid-
summer short-winged, non-diapause group, except that G. lacustris populations
also include overwintering brachypters.

The first fifth instar larva was captured on 12 June, and the first new, non-
diapause adults (both brachypters and macropters) on 26 June. The last over-
wintered imago (one brachypterous female) was caught on 10 July.

The last young, definitely non-diapause adult (a brachypterous female),
incompletely pigmented and semi-soft, with the largest oocytes at least 860 um,
was captured on 10 July. However, still on 24 July one brachypterous female, soft
and incompletely pigmented (signs of darkening venter could be traced), still had
oocytes of 550 um maximum length. Furthermore, on the same day one semi-soft
and incompletely pigmented brachypterous female (with signs of dark venter) with
maximal oocyte length of 480 um was caught. These two oocyte measurements lay
at the lower boundary of the class “maturing” of Andersen (1973: 500 um) but

Fig. 2. Seasonal changes in the proportions of the different female forms of Gerris lacustris. Symbols as
in fig. 1 but instead of dark (light) venter read dark (light) thoracic venter. The diagram is an
approximation drawn on basis of 60.6% of the brachypter and 42.2% of the macropter females.

10 42 28 100 1724 44 49 65 27 32
%
80;
60:
40:
20;

D EEE Il

16 16 30 12 26 10 24 21 4

APRIL M AY JUNE JULY AUGUST

VEPSÄLÄINEN & NIESER: Life cycles of Dutch Gerris species 209

considerably exceed the 240 um used by Vepsäläinen (1974a) in routine scrutiny. It
is not clear whether these two females really were diapausing or not. The
coloration of the thoracic venter indicates diapause (accordingly a later resorption
of oocytes is to be expected). The colour of the venter is, however, only a good
working guide, as in the same samples there is one semi-hard, ventrally dark (only
sternites 6 and 7 medio-apically light) brachypterous female with eggs. In this case
there could not be any question of an overwintered adult as the ventral coloration
was still partially brown-black (a sign of ongoing pigmentation). Another puzzling
individual is the macropterous female caught on 7 August, which was incompletely
pigmented, semi-soft/semi-hard and had only one, 365 um long oocyte in its
otherwise undeveloped ovaries.

Thus it is clear that whether the females were classified as maturing by the
criterium of minimum oocyte length of 240 um or 500 um, it is preferable to check
for additional signs of diapause in border cases as well. The positive correlation
between reproductive activities and poorly developed adipose tissue, and diapause
and well-developed adipose tissue seems to hold true in individuals which have
attained at least semi-hard chitin, but is poorly developed in soft and semi-soft
imagoes, even if completely pigmented.

Non-diapause adults were still found in late August and early September,
although the numbers were low. The first diapause imago occurred on 10 July, and
on 24 July diapause imagoes dominated in the total material.

The positive relation between a light thoracic venter and a non-diapause state in
G. lacustris is good in our Dutch material. Only one possible exception (above) was
found. Moreover, at least four macropterous females were seen (on 24 July), in
which the general impression was all-dark. A closer inspection revealed that in all
these individuals the pattern of lighter non-diapause adults could be seen. The
coloration was only more grey-brown than usually (“‘smoky’’), but still contrasted
against the black areas of the thoracic venter. The coloration was much the same
as in the non-diapause type of G. paludum F. in fig. 3 by Vepsalainen (1974a).

Brachypterous G. lacustris adults have undeveloped flight muscles. Diapause
macropters develop functional muscles, which, after overwintering, are histolysed.
In our Dutch material we already have macropterous females with histolysing
muscles on 16 May. In mid summer, diapause imagoes seem to reach flight ability
in about two weeks or shorter: two females collected on 24 July were semi-hard,
totally pigmented, with mesoalinotum II and the diameter of indirect flight muscle
fibres 30-50 um and 32-45 um, respectively.

Sex ratio of brachypters is perhaps female-weighted (43% males, n=174,
P=.07); in macropters the male percentage is 51 (n=475).

Other species

Altogether 23 G. gibbifer Schumm. adults (10 G, 13 ®) were collected, all
macropters (one female with hemielytra 7.8 and hind wings 7.0-7.1). The first new
generation adult was caught on 26 June; it was a semi-soft female with
undeveloped oocytes. Oocyte maturation could not be detected in an already

210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 7, 1977

semi-hard/hard female on 10 July. Thus the voltinism of Dutch G. gibbifer cannot
be determined on the basis of our material.

Only two observations of overwintered G. rufoscutellatus Lat., probably the same
female, were made: on 30 May, and 12 June, when its abdomen was swollen with
eggs. No larvae were found.

Midsummer colonization flights

Usually little can be said about colonization flights without marking-recapture
work. The temporary drying up of pond N13 (dry on 26 June, water on 10 July),
however, gave some information.

On 26 June the bottom and the bordering vegetation were checked for Gerris:
none were found. On 10 July one G. thoracicus macropterous male and one
G. lacustris macropterous female (semi-hard, light thoracic venter, eggs; thus a
midsummer, non-diapause adult) were collected. On 24 July another G. thoracicus
macropterous male and G. lacustris macropterous female (semi-hard, light thoracic
venter, eggs, totally pigmented mesoalinotum and well developed indirect flight
muscles) were taken.

On 7 August two fifth instar larvae of G. gibbifer and three G. thoracicus females
were netted. The G. gibbifer larvae may have developed from eggs which survived
the drought, as on 30 May and 12 June there was a G. gibbifer female in the pond.
Of the G. thoracicus females, two had eggs and the third was in diapause. As no G.
thoracicus larvae were collected from N13, it is improbable that the adults had
developed there.

Accordingly, colonization flights of mid summer reproducing G. lacustris and G.
thoracicus do occur. Earlier straight evidence is rare. Fernando (1959) reports one
immigrant G. odontogaster female which produced offspring. Here the female was
fertilised before the migration, as perhaps, were some females of our immigrants
to population N13. There seems to be a clear difference therefore in the stage of
maturation between colonising overwintered and colonizing midsummer females,
as spring migrants have undeveloped or very poorly maturing oocytes (Landin &
Vepsalainen, 1977).

DISCUSSION

The life cycles and wing lengths of the studied Gerris species are by and large
similar to those in Poland (Vepsalainen & Krajewski, 1974) and Hungary
(Vepsalainen, 1974a). The Dutch populations differ, however, in these respects
from more northern ones in Finland (see Vepsäläinen, 1974b). Admittedly, there
are more non-diapause macropters in the mid-summer generation in Hungary
(and, considering G. lacustris, also in Poland). This could reflect the higher degree
of temporariness of more southern and continental pond habitats.

The diapause of Dutch Gerris seems to be fixed at about the same date, and
presumably by the same mechanism, as that of Finnish, Polish and Hungarian
populations (for a generalized model, see Vepsäläinen, 1974c: fig. 2).

In northern Europe early winter understandably favours diapause determination

VEPSALAINEN & NIESER: Life cycles of Dutch Gerris species 211

by decremental changes of day length. On the average the potential reproductive
season is longer and the risks of drying up of population sites are smaller in
Atlantic climates than in continental ones at the same latitudes. Thus it could be
assumed that in Dutch populations selection would operate towards a later
termination of reproductive activities. This could be achieved in at least two ways:
by later onset of diapause and/or longer life span of the midsummer, non-diapause
adults.

Usually the diapause of insects is determined by the absolute lenghts of daily
illumination period (Danilevskii, 1965) but in Gerris the change of length is critical:
shortening day lengths determine the diapause. Thus it is not easy to postpone the
onset in diapause in Gerris. The associated genetic changes would have to be
radical, and would be possible only in effectively isolated populations under strong
selective pressure. The Dutch populations in this respect are not submitted to
strong natural selection and are insufficiently isolated. From the degree of tempor-
ariness of our study ponds it can be presumed that gene flow between local Gerris
populations is notable, which is in agreement with the general views on migration
strategies of the denizens of more or less temporary habitats (e.g. Southwood,
1962; Dingle, 1974; Vepsalainen, 1974b). Even long range flights are known
(Leston, 1956). These tend to swamp the effects of local selection pressures.
Moreover, a recent simulation study on Gerris populations (Jarvinen & Vepsa-
läinen, unpublished) showed that even in the absence of imminent threat of
unfavourable conditions it may sometimes (e.g. in habitats of low productivity) be
advantageous to limit the number of successive generations.

The strategy to postpone the end of reproductive activities by cutting down the
mortality of the midsummer non-diapause adults is likely to work in Gerris.
Unfortunately we lack Dutch data on last first-instar larvae in autumn. Con-
sequently, nothing can be said of the approximate last day of egg-laying of Dutch
Gerris. Here would be an important field of study: to work out the length of the
reproductive season in terms of last egg-laying dates of Gerris species in different
parts and climates of the western Palaearctic. The theoretical value of such a study
is evident, as the onset of diapause seems to occur about the same date
everywhere irrespective of the widely differing lengths of the potential reproduc-
tive season.

Another point of discussion comes from our observation of two completely
pigmented new-generation G. odontogaster females (semi-hard and semi-soft) with
dark venter and undeveloped oocytes, captured on 12 June. The observed features
together are a reliable sign of diapause. Vepsäläinen (1974d) obtained such
laboratory offspring of Finnish G. odontogaster which were reared in short,
lengthening illumination. The applied laboratory days were shorter than Finnish
late instar larvae can meet in nature in early summer. Such females are expected
to be very rare in natural populations, as the selection pressures are heavy against
diapause behaviour in early summer. It cannot, however, be ruled out that oocyte
maturation both in the Finnish laboratory reared and the Dutch females was only
retarded and would have started later in somewhat longer days, producing
exceptionally new-generation females with eggs, though with dark venter. Such
individuals have very rarely been observed (e.g. Vepsalainen & Krajewski, 1974, in

2112. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 7, 1977

G. lacustris). We presume that retarded maturation instead of real diapause is the
more likely explanation in some individuals which develop in short, lengthening
days — i.e. exceptionally early in summer — in nature.

ACKNOWLEDGEMENTS

Thanks are due to Hilkka Vepsäläinen who made the life history diagrams, Olli
Järvinen who made valuable suggestions to the text, and Elisabeth de Groot-Taat
who checked the English. Finally thanks are due to the S.B.B. (State Forest
Service), section Gelderland, for permission to sample in closed or otherwise
protected areas.

REFERENCES

Andersen, N. M., 1973. Seasonal polymorphism and developmental changes in organs of flight and re-
production in bivoltine pondskaters (Hem. Gerridae). — Ent. Scand. 4: 1—20, 11 figs.
Danilevskii, A. S., 1965. Photoperiodism and seasonal development of insects: ix + 283, 62 figs. —

London.

Dingle, H., 1974. The experimental analysis of migration and life-history strategies in insects. In: L. B.
Browne, Experimental analysis of insect behaviour: 329—342, 4 figs. — Berlin.

Fernando, C. H., 1959. The colonization of small freshwater habitats by aquatic insects. 2. Hemiptera
(the water-bugs). — Ceylon J. Sci. (Bio. Sc.) 2(1): 5—32, 7 figs.

Jarvinen, O., 1976. Migration, extinction and alary morphism in water-striders (Gerris F.). — Ann.
Acad. Sci. Fennicae (A IV) 206: 1—7.

Krajewski, S., 1969. Pluskwiaki wodne (Heteroptera) rzeki Grabi i jej terenu zalewowego. — Polsk. Pis-
mo. Ent. 39: 465—513.

Landin, J., & K. Vepsäläinen, 1977. Spring dispersal flights of pond-skaters Gerris spp. (Heteroptera).

; — Oikos 29: 156—160, 1 fig.

Leston, D., 1956. The status of the pondskater Limnoporus rufoscutellatus in Britain. — Ent. Month.
Mag. 92: 189—193.

Poisson, R., 1957. Hétéroptères aquatiques. — Faune de France 61, 263 pp., 185 figs.

Southwood, T. R. E., 1962. Migration of terrestrial arthropods in relation to habitat. — Biol. Rev. 37:
171—214, 1 fig.

Vepsalainen, K., 1971. The role of gradually changing daylength in determination of wing length, alary
dimorphism and diapause in a Gerris odontogaster (Zett.) population (Gerridae, Heteroptera)
in South Finland. — Ann. Acad. Sci. Fennicae (A IV) 183: 1—25, 7 figs.

—., K., 1973. The distribution and habitats of Gerris Fabr. species (Heteroptera, Gerridae) in Fin-
land. — Ann. Zool. Fennici 10: 419—444, 10 figs.

——, 1974a. The wing lengths, reproductive stages and habitats of Hungarian Gerris Fabr, species
(Heteroptera, Gerridae). — Ann. Acad. Sci. Fennicae (A IV) 202: 1—18, 6 figs.

———, 1974b. The life cycles and wing lengths of Finnish Gerris Fabr. species (Heteroptera, Gerri-
dae). — Acta Zool. Fennica 141: 1—73, 14 figs.

— ., 1974c. Determination of wing length and diapause in water-striders (Gerris Fabr., Heteropte-
ra). — Hereditas 77: 163—176, 2 figs.

——, 1974d. Lengthening of illumination period as a factor in averting diapause. — Nature (Lond.)
247: 385—386, 2 figs.

Vepsäläinen, K., & S. Krajewski, 1974. The life cycle and alary dimorphism of Gerris lacustris (L.) (He-
teroptera, Gerridae) in Poland. — Not. Ent. 54: 85—89, 2 figs.

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MUS. COMP. ZOOL.
LIBRARY
DEEL 120 AFLEVERING 8 1977

MAR 8 1978

HARVARD
UNIVERSITY

TIJDSCHRIFT
VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

F. WILLEMSE. — Classification and distribution of the Sexavae of the Melanesian
Subregion (Orthoptera, Tettigonioidea, Mecopodinae), p. 213—277, text-figs.
1—7, pls. 1—18.

| Tijdschrift voor Entomologie, deel 120, afl. 8 Gepubliceerd 30-XII-1977 |

CLASSIFICATION AND DISTRIBUTION OF THE SEXAVAE
OF THE MELANESIAN SUBREGION (ORTHOPTERA,
TETTIGONIOIDEA, MECOPODINAE)

by

FER WILLEMSE

Laurastraat 67, Eygelshoven, The Netherlands
With 7 text-figures and 18 plates

ABSTRACT

An attempt is made to classify the species of the genera Sexava Walker, Segestes Stäl and Segestidea
Bolivar on the basis of previously not used characters, such as the venation of the fore wing and the
morphology of the male stridulatory apparatus.

The following new synonyms and new combinations are proposed: Eumossula C. Willemse, 1957 =
Segestidea I. Bolivar, 1903; Segestidea gracilis (C. Willemse, 1957), comb. nov. (from Eumossula);
Segestes grandis C. Willemse, 1955 = Sexava coriacea (Linné, 1758); Sexava femorata C. Willemse, 1940
= Segestidea princeps 1. Bolivar, 1903 = Segestidea novaeguineae (Brancsik, 1897), comb. nov. (from
Moristus); Segestidea hanoverana C. Willemse, 1957 = Segestidea leefmansi (C. Willemse, 1940), comb.
nov. (from Sexava); Segestidea insulana C. Willemse, 1957 = Segestidea uniformis (C. Willemse, 1940),
comb. nov. (from Sexava); Segestidea acuminata (Kastner, 1934), comb. nov. (from Segestes); Segestidea
rufipalpis (C. Willemse, 1966), comb. nov. (from Sexava). Besides the following new taxa are described:
Segestes stibicki sp. n., Segestes cornelii sp.n., Segestes brevipennis sp.n., Segestidea gracilis simulatrix
ssp.n. and Segestidea marmorata occidentalis ssp.n.

Keys to the taxa and distribution maps are given. The range of the species and subspecies is now in
agreement with the general zoogeography of the area concerned; previous records proved to be partly
erroneous.

CONTENTS
Introduction: u...54bere ren Pe ie gies ne AE at GLEN ee Sant a bags se ee 213
Meateriallandiacknowledgementsereu ee RISI 214
Geneniciand specificicharactersimiSexavac: PAPE TEE 215
Wey tothe general ur we RAT NE E ORO MIRTO I ENEN 218
Sexava Walker, Fb Sth cit MD eR TE RENTE Se My Ly Ne 218
Keyatosthespecies of Sexavart rue. NRE yi O ee: 219
Segestes;Ställ- iu eee din geeks sity der ee NE hay oy wee at. ok cen tahitian dE 227
Key to the species of Segestes from New Guinea and Bismarck Archipelago .......... _230
Segestidea Bolivar: cin. INA RL EAD DART O N TD 240
KeyitomheispeciesiofSegestide a Ne N. 241
RÉfETENCES ARMES LME RR det. 256
Ind RARES CUT EAA TR A ET RE ER IRE EU SONO ER AT ANNEE Re MURS NN 277
INTRODUCTION

The species of the genera group Sexava Walker, Segestes Stal and Segestina I.
Bolivar are large katydids, called ““coconut treehoppers” by entomologists. Some

218

214 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

of them have been reported as important pests of coconut trees in the Moluccas,
New Guinea and the Bismarck Archipelago.

Dr. J. L. Stibick of the Department of Agriculture, Stock and Fisheries,
Konedobu, Papua, has sent me material for identification since 1971. It came out
that the taxonomy of this group of genera was considerably confused and generic
distinction was not well understood. As a result, some species were described
twice, in different genera.

The range of the species of the Sexavae covers the Philippines, Celebes, the
Moluccas, New Guinea and the Bismarck Archipelago. The present study deals
mainly with species occurring in the Melanesian Subregion.

Only the more important taxonomic references are mentioned. A more
extensive, although incomplete, bibliography can be found in Beier (1966: 311 —
316). The measurements given throughout the text concern the lengths and are in
millimeters. The nomenclature of the wing venation is that given by Ragge (1955).
Figures of the male stridulatory apparatus and abdominal terminalia of the taxa of
each genus are nearly on the same scale, those of the whole insects not. On the
distribution maps, only reliable localities are indicated, while a few previous
records are omitted as no material was at hand and identification appears
doubtful.

MATERIAL AND ACKNOWLEDGEMENTS

Depositories of types and other material are given in abbreviated form
throughout the text, as follows:

ANSP Academy of Natural Sciences, Philadelphia;

BMNH British Museum (Natural History), London;

BPBM Bernice P. Bishop Museum, Honolulu;

CAS California Academy of Sciences, San Francisco;

CW Willemse’s collection, partly Natuurhistorisch Museum, Maastricht,
partly author’s address;

DASF Department of Agriculture, Stock and Fisheries, Konedobu, Papua;

IEM Instituto Espanol de Entomologyéa, Madrid;

ITZ Instituut voor Taxonomische Zoölogie, Amsterdam;

NMB Naturhistorisches Museum, Basel;

NMW Naturhistorisches Museum, Wien;

NR Naturhistoriska Riksmuseet, Stockholm;

NS Naturkundemuseum, Stettin;

RNH Rijksmuseum van Natuurlijke Historie, Leiden.

My thanks are due to the following persons for helping me with material and
valuable information: C. Baroni Urbani, Basel; P. H. van Doesburg, Leiden: J. L.
Gressitt, Honolulu; A. Kaltenbach, Vienna; the Keeper of Entomology of the
British Museum (Natural History), Mrs. L. M. Pitkin and D. Ragge, London; T.
Kronestedt, Stockholm; G. Kruseman, Amsterdam; V. Llorente, Madrid; D.
Rentz, San Francisco; H. Steinmann, Budapest; J. N. L. Stibick, Konedobu.

WILLEMSE: Sexavae of Melanesian Subregion 215

GENERIC AND SPECIFIC CHARACTERS IN SEXAVAE

The Sexavae, as understood by C. Willemse (1961: 93), refer to the ‘‘Sexava-
Gruppe” of Karny (1924: 144) and Kästner (1934: 24) and comprise the genera:
Sexava Walker, 1870, Segestes Stäl, 1877, and Segestidea I. Bolyevar, 1903.
Previously (Karny, 1924: 143; Kästner, 1934: 24), the name Sexavae was used in a
much wider sense, covering also the Phrictae and Mossulae groups of genera.
These three groups of genera are united under one of the two tribes recognized in
the subfamily Mecopodinae, the Sexavini (called Moristini by Redtenbacher,
1892: 189; Segestini by Hebard, 1922: 176; Sexavae by Karny, 1924: 143, and
Kästner, 1934: 24; Sexavinae by C. Willemse, 1961: 93; Sexavini by Beier, 1966:
305). Keys to the tribes and genera were given by Redtenbacher (1892: 189),
Caudell (1916: 2), Karny (1924: 143, 147), Kästner (1934: 24, 32) and C. Willemse
(1961: 93, 94, 107, 111). While the distinction between the Phrictae and the
remaining genera of the tribe appears clear, that between the Mossulae and the
Sexavae is more gradual and demands further study.

Distinction between the three genera of the Sexavae was based on the presence
or absence of a posterior dorso-apical spine of the fore and mid tibiae, and the
comparative measurements of body, fore wing, head and pronotum (Caudell, 1916:
3; C. Willemse, 1957: 42; 1961: 107). In Sexava and Segestidea, the apex of the fore
and mid tibiae bears, at least at the posterior side, a small dorsal spine. This spine
is not to be confused with one or more spines located more proximally on the
dorsal margins of the tibiae. In Segestes, the dorso-apical spines are believed to be
lacking. A study of the available material reveals that this character is indeed a
reliable one, but that there are exceptions. In three out of 38 specimens assigned
to Segestes decoratus Redtenbacher, dorso-apical spines of fore and mid tibiae are
present. In these specimens the spines are sometimes very small and not uniformly
present on all fore and mid tibiae. An explanation of these exceptions can be
found in the remarkable stability of the occurrence of the dorso-apical spines in
the subfamily. Actually, the presence of these spines has been used as a subfamily
character as early as Brunner von Wattenwyl (1878: 10) until recently (Beier, 1955:
246; 1962: 2), although Redtenbacher (1892: 183) already gave comments on this
point and re-defined the Mecopodinae. A renewed study of other Segestes
material at hand, i.e. the remaining part of the material in the Vienna museum
(Kaltenbach, in litt. 11.11.1976) and of the type-species vittaticeps Stäl, proves that
the dorso-apical spines in that material are indeed lacking. Therefore it seems
reasonable, in spite of the very few exceptions, to consider the presence or
absence of dorso-apical spines of the fore and mid tibiae a character of generic
significance.

The distinction between Sexava and Segestidea, as indicated previously, was
arbitrary (Caudell, 1916: 3; C. Willemse, 1957: 42; 1961: 107). Among the
characters mentioned, it was noted that the fore wing in Sexava is comparatively
wider than in Segestidea, the venation not being mentioned. Also Redtenbacher, in
his study on the wing venation (1886: 179, pl. 11 fig. 24), discussed the wings in
Moristus (=Sexava), but omitted the true nature of its typical venation.

Comparison of the male fore wings of Sexava, Segestidea and Segestes, reveals

216 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

R SE È

Aa
7 4

Cu2

Fig. 1. Segestidea, right fore tibia and tarsus, posterior view. Figs. 2—3. Left male fore wing: 2, Sexava
coriacea (L.); 3, Segestidea novaeguineae (Brancsik).

that the fore wings overlap each other basally over a short distance in Segestidea
and Segestes, whereas in Sexava the overlap is conspicuous over most of their
length. It proves that the course of the first cubital vein, Cul (which may fuse or
not with the variably developed MP), in Sexava is remarkably different from that
in both other genera. This character affects the stridulatory area of the male fore
wing and is considered more reliable for a natural classification of species at
generic level than previously used characters.

The flexed fore wing in the species of Sexavae is folded along Cul. The result of
this fold is to bring the areas behind Cul (cubital and anal areas) into a roughly
horizontal position over the abdomen, the remainder of the wing being roughly
vertical. In Segestidea and Segestes, Cul and its fold run obliquely to the hind
margin of the wing, almost reaching the latter at a short distance from the wing-
base (text-figs. 3, 6, 7). From there, Cul may more or less merge with the
archedictyon or continue parallel but very close to the hind margin of the wing.
Cul and its fold delimite a sharply defined triangular area at the wing-base. This
area contains, in the male, the most important parts of the stridulatory organ: the
file (=Cu2) of the left and the mirror and its frame of the right fore wing (text-figs.
4—7). In Sexava, however, Cul and its fold reach the hind margin of the fore wing
considerably more distally, near or beyond the middle of the wing length and do
not delimite a sharply defined triangular area (text-figs. 2, 4, 5). As a result, the
first cubital area (between Cul and Cu2) in Sexava is conspicuously wide and

WILLEMSE: Sexavae of Melanesian Subregion 217

elongate, which is demonstrated also by the large overlap of the flexed wings, and
the considerable extension of the hind margin of the flexed wing beyond the
median line of the abdomen. It is noted here that these characters are very distinct
in the male, but far less so in the female.

The high taxonomic significance of sound produced by Tettigoniids is widely
recognized. Nevertheless, the structure of the stridulatory organ has been little
used as a character. The present study makes use of some attributes of the male
stridulatory organ, which can be easily examined: the file of the left fore wing and
the mirror and its frame of the right one. These have been studied in most of the
available males. The length, width and shape of the file, the number and spacial
arrangement of its teeth and the shape, and especially the frame of the mirror of
the opposite wing, are in most species and subspecies slightly variable and specific
distinction is sometimes conspicuous. I did not have the opportunity to analyse the
characters of the file as extensively as has been done in some Phaneropterine
genera (Emsley, et al.,1967, 1969; Ragge, 1969; Huxley, 1970; Emsley, 1970; Moss,
et al.,1970). Instead, I give a short description together with figures of the file and
the mirror. As to the latter, it appears that the frame of the mirror, especially a
vein modified into a fold along the hind margin of the mirror, is important for
specific recognition.

Stridulation of the female in the Mecopodinae has been recorded by Karny
(1924: 143), but his record refers to Macrolyristes, which is a member of the tribe

Figs. 4—7. Base of left and right male fore wing, dorsal view: 4—5, Sexava coriacea (L.); 6—7,
Segestidea novaeguineae (Brancsik).

218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

Mecopodini. Recently, Lloyd & Gurney (1975: 47—50) reported labral stridulation
of a female attributed to Sexava femorata (= Segestidea novaeguineae). However,
the morphological substrate of this kind of stridulation (labrum versus mandibulae)
is obscure. A closer study of the fore wings of the female in Sexavae reveals that
there is a stridulatory apparatus. It consists of rows of minute spines located on the
dorsal surface of the veins near the hind margin of the cubito-anal areas of the
right fore wing, where the hind margin of the left wing may function as a scraper.
Due to uniform appearance throughout the species of the group, no reliable
characters were found which would associate females with males of the same
species. Similar spines were often found on the male right fore wing, just distally of
the mirror.

Characters found to be more or less reliable at species or subspecies level, may
be summarized: general appearance; length of fastigium of vertex; shape of
pronotal lateral lobe, convexity of pronotal dorsum; fore wing (length, width,
shape, archedictyon, apex, male stridulatory apparatus); spines of legs; shape of
male subgenital plate and cercus; length of ovipositor; coloration of head, thorax,
fore wing and legs.

Key to the genera Sexava, Segestes and Segestidea

1. Fore and mid tibiae with a small, posterior, dorso-apical spine (text-fig. 1)
(refers to tip of tibia only, not to dorsal margins; if tibia stretched forward,

posterior.is “lateral? side), it La 2
— These spines lacking (with very few exceptions in Segestes decoratus); venation
of fore wing as in Segestidea, see below ................ Segestes Stal

2. Cul and similarly located fold of fore wing running obliquely toward hind
margin of wing at short distance from wing-base, delimiting well-defined
triangular area, in male containing the main attributes of the stridulatory organ
(text-figs. 3, 6, 7); flexed fore wings, except for triangular area at base,
narrowly overlapping each other, hind margin not widely extending over
medianilinciofiab dome eee e Segestidea I. Bolivar

— Cul and its fold running to a point of the hind margin of fore wing near or
beyond middle of wing-length, without delimiting a well-defined triangular
area (text-figs. 2, 4, 5); flexed fore wings overlapping each other widely and
over great distance of wing-length, hind margin extending far beyond median
line of abdomen over almost whole length ............ Sexava Walker

Sexava Walker, 1870

Sexava Walker, 1870: 437.
Moristus Stäl, 1873: 47 (type-species, by monotypy: Gryllus (Tettigonia) coriacea Linné, 1758)

Type-species, by monotypy: Gryllus (Tettigonia) lanceolata Stoll, 1813.

Walker based his genus on a female from Ceram, assigned by him with some
doubt to Stoll’s lanceolata. This species is synonymous with Linne’s coriacea.
Walker’s specimen, however, does not agree with /anceolata (= coriacea), but with
Stäl’s nubila, described in 1874, after Walker’s publication. Stability and uniformity

WILLEMSE: Sexavae of Melanesian Subregion 219

of nomenclature is best served when neglecting Walker’s misidentification (Article
70 (a) (iii) of the International Code of Zoological Nomenclature). Therefore I
propose to maintain the current type-species.

Sexava, as indicated by the key above, is a natural group of species,
characterized by the presence of posterior dorso-apical spines of fore and mid
tibiae (text-fig. 1) in combination with the unusual course of the first cubital vein
(Cul) of the fore wing (text-figs. 2, 4, 5), resulting into the large overlap of the
flexed fore wings both with each other and with the median line of the abdomen.

Up till now, the following taxa have been arranged under Sexava:
coriacea (Linné, 1758) (= lanceolata Stoll, 1813)
nubila (Stal, 1874) (Moristus)
novaeguineae (Brancsik, 1897) (Moristus)
karnyi Leefmans, 1927
femorata C. Willemse, 1940
uniformis C. Willemse, 1940
leefmansi C. Willemse, 1940
grandis (C. Willemse, 1955) (Segestes)
rufipalpis C. Willemse, 1966

Of this list, five taxa apparently belong to Segestidea, while only four taxa fit
Sexava in its present concept: coriacea, nubila, karnyi, and grandis. The last will be
synonymized with coriacea in the present paper.

Key to the species of Sexava

l. d, 9: smaller, length of fore wing 50—60 mm; lower and part of inner side of
proximal half of hind femur solid black; &: cercus with short apex (pl. 3 fig.
22), teeth of stridulatory file in posterior half very widely spaced (pl. 4 fig. 28)
(Togian Is. and opposite part of Celebes) ............ karnyi Leefmans

— 6, 2: larger, wing-length more than 65 mm; hind femur without black parts;
dg: cercus with apex longer (pl. 3 figs. 18—21); teeth of stridulatory file more
closelyset(pls4. figs; BO) keke ie ea wy ee ER ED ARS. 2

2. &: subgenital plate wider, apical incision shorter, lobes wider, styli shorter (pl.
3 figs. 12—14); cercus with longer, more attenuate apex (pl. 3 figs. 18—19);
teeth of stridulatory file narrowly set (pl. 4 figs. 23—25); g, 9: pronotal lateral
lobe usually as long as high; apex of flexed wings reaching middle third of hind
tibia; 9: subgenital plate usualiy shorter than wide; ovipositor longer, 40—50
mm, apex of flexed wings usually not reaching tip of the latter (Sangihe Is.;
central Moluccas; ? Celebes; ?? New Guinea) .......... coriacea (Linné)

— d: subgenital plate narrower, apical incision longer, lobes narrower, styli
longer (pl. 3 figs. 15—16); cercus with shorter, less attenuate apex (pl. 3 figs.
20—21); teeth of stridulatory file coarser (pl. 4 figs. 26—27); 3, 9: pronotal
lateral lobe usually narrower, being shorter than high; apex of flexed wings
reaching beyond middle of hind tibia, usually apical third; 9: subgenital plate
usually as long as wide: ovipositor shorter, 30—43 mm, apex of flexed wings
reaching at least tip of the latter (Northern Celebes; Talaud and Nanusa Is;

220 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

southeastern Moluccas: Kei and Aru Is.; western and northeastern New
Guinea)TF. Meelis donnaient abbienti: nubila (Stal)

Sexava coriacea (Linné, 1758)
(text-figs. 2, 4, 5, pl. 1 figs. 1—4, pl. 3 figs. 12—14, 18, 19, pl. 4 figs. 23—25,
pl. 5 fig. 29, map 1)

Gryllus (Tettigonia) coriaceus Linné, 1758: 430.

Gryllus (Tettigonia) lanceolata Stoll, 1813: 23, pl. 10a figs. 39-40.
Locusta lanceolata; de Haan, 1843: 214 (partim).

Moristus coriaceus; Stal, 1873: 95.

Sexava coriacea; Kirby, 1906: 359.

Segestes grandis C. Willemse, 1955: 36, fig. 2. syn. nov.

Sexava grandis; C. Willemse, 1957: 38 (footnote).

Material studied: Soela Mangoli, Pasi Spah, 18—19.iii.1930, Snellius Exp. (1&
RNH); Ins. Buru, H. Kühn, coll. Br. v. W. (14 NMW); Batjan, Staudinger, coll.
Br. v. W. (13 NMW); Ternate, 1894, W. Kükenthal (14 NMW); Halmaheira (1 ¢
19 CW), 1894, W. Kükenthal (19 NMW), Djailolo forêt Todowangi, 16.ii.1929,
Prince Léopold (14 CW); Sangir, A. Reyne, ex coconut palms (14 19 BMNH);
Sangir, xi.1948, C. Franssen, coconut leaf, ex coll. S. Leefmans (29 ITZ); Ambon
(lg CW, 14 12 ITZ), 1863, Hoedt (19 RNH), 1864, Hoedt (6g 29 RNH),
Staudinger, coll. Br. v. W. (14 NMW), 1933, coll. R. Ebner, Rehn don. (19
NMW), 1859, Doleschal (19 NMW), 30 m, 1.ix.1961, A. M. R. Wegner, at light
(19 CAS), Waai, 1.viii.1966, A. M. R. Wegner (1 9 CAS); Molukken, Depuiset,
coll. Br. v. W. (19 NMW); Java (1g 19, De Haan vidit, RNH); Nieuw Guinea
(18 CW); Celebes, 1884, Musschenbroek (29 ITZ); Obi I. (19 BMNH).

Segestes grandis: type-series, & holotype: Obi Island, Anggai 30.v.1953; ©
paratype: Obi Island, Laiwui, 20.ix.1953 (both specimens with appropriate
identification and type-labels) (CW).

Additional material: Obi (14 CW), Telaga, 7.viii.1953 (19 CW), Laiwui,
29.ix.1953 (23 19 CW, 23 29 RNH), Wajaloar — Obi, 27.viii.1953 (14 RNH)
(all identified by C. Willemse).

The Linnean type of Gryllus (Tettigonia) coriaceus could not be traced in London
(Ragge & Mrs. Pitkins, in litt. 5.iii.1976).

The possible synonymy of Stoll’s lanceolata with Linné’s coriacea is discussed
extensively by Karny (1931: 78-82). Stoll’s material was from the collection of J.
Raije van Breukelerwaert. This collection was sold 3 July 1827 in Amsterdam and
the name lanceolata is found under the numbers 372 and 374 (p. 76) of the
“Catalogue ... d’objets d’histoire naturelle... Joan Raye de Breukelerwaert...
sera vendu...a Amsterdam ...’’. However, the specimens in question could not
be traced in the collections at Amsterdam (ITZ) or Leiden (RNH). Nevertheless,
the length of the female wings and ovipositor of Stoll’s figure 39 makes
identification possible. That figure perfectly fits coriacea and not nubila. Therefore
I again propose to synonymize lanceolata with coriacea.

WILLEMSE: Sexavae of Melanesian Subregion 221

In the original description, Segestes grandis C. Willemse, 1955, was clearly
distinguished from other members of that genus. In his 1957 paper, C. Willemse
allocated the species correctly to Sexava, but distinction among species of that
genus was omitted. Comparison of the types and additional topotypes of grandis
with material of coriacea clearly reveals that grandis represents merely large
individuals which are within the range of variation of coriacea. I propose to
synonymize both taxa.

Sexava coriacea is defined as indicated in the key. A general description can be
found under Segestes grandis by C. Willemse (1955: 36, fig. 2). Reliable characters
are the male subgenital plate (pl. 3 figs. 12—14), cercus (pl. 3 figs. 18—19) and
stridulatory file (pl. 4 figs. 23—25). The latter is slightly fusiform and arcuate, 4—8
mm long, number of teeth 48—92, the anterior 25—45 ones almost blunt and
covering the anterior fourth, the other teeth sharp and arranged over the
remaining length of the file. Width of the file increasing in the anterior third,
reaching maximum 0.5—0.9 mm in the middle third, decreasing again posteriorly
to about half maximum width. Spacing of teeth narrow, distinctly increasing from
the anterior to posterior end of the file, between the posterior teeth about 2—4
times as large as between the anterior sharp teeth. The number of teeth and the
maximum width depend mainly on the length of the file, while the spacing of the
teeth is quite uniform. The mirror (pl. 5 fig. 29) is about as long as wide, roughly
circular with the anterior margin almost straight. Fold of the mirror strongly
inflated in the middle, slightly extending over the mirror, its outline from almost
straight to slightly sinuate.

Variation. The measurements, especially the length and width of the fore wing
(pl. 1 figs. 1—4) vary considerably. While the ovipositor usually extends beyond the
apex of the flexed wings, they may be of the same length in the females from
Sangihe and Obi Islands (pl. 1 figs. 3—4). The stridulatory file of the studied males
from Buru and Ambon (pl. 4 fig. 25) is shorter (4—5.3 mm) and the number of
teeth smaller (47—72), while the file in males from other islands (pl. 4 figs. 23, 24)
is longer (5.4—8.2 mm) and the number of teeth larger (72—95). The male
subgenital plate is variable (pl. 3 figs. 12—14) in width, the margins of the apical
incision vary between smooth and slightly serrate and the tips of the lobes from
truncate to slightly emarginate. Variation is also observed in the female subgenital
plate. As in the stridulatory file, the variation of the subgenital plate of both sexes
corresponds with the area of occurrence (compare also Leefmans, 1927b: pl. 5).
However, the shape of the male cercus is quite uniform (pl. 3 figs. 18, .19). The
pronotal lateral lobe of some specimens is slightly shorter than high. The general
coloù, usually green, is sometimes brown. Fore wing and hind femur are always of
general colour, lacking any pattern.

Measurements: body & 60—85, © 60—71; fore wing & 65—80, 9 75—86; hind
femur g 43—48, 9 45—51; ovipositor 40—50.

Distribution. The range covers the central islands of the Moluccas and the
Sangihe Is. The occurrence in Celebes and New Guinea needs confirmation (map
1).

Localities: Ambon (Redtenbacher, 1892: 201; Krauss, 1903: 747; Hebard, 1922:
180; Karny, 1924: 151; Leefmans, 1927b: 13); Buru (Karny, 1926: 183); Batjan

222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

(Karny, 1924: 151); Sula Is. (Karny, 1924: 151), Pasi Spah near Sula Mangole; Obi
(Hebard, 1922: 180; Karny in Leefmans, 1927b: 13), Anggai and Laiwui (C.
Willemse, 1955: 36), Wajaloar and Telaga; Banggai Is., Labobo (Leefmans, 1927b:
13); Ternate (Brunner v. W., 1898: 199; Leefmans, 1927b: 13); Halmahera, Tobelo
(Leefmans, 1927b: 13), Djailolo (C. Willemse, 1933: 9); Sangihe Is. (Leefmans,
1927b: 13); ? Celebes (Karny, 1924: 151).

Discussion. The species is well-defined in the male. However, distinction
between the female of coriacea (pl. | figs. 3, 4) and the form of nubila (pl. 1 fig. 6),
which occurs in the Talaud and Nanusa Is., may be very doubtful. Sometimes the
following characters are helpful: the wider thorax, the absence of a series of
yellowish dots on the radial area of the fore wing, and the longer ovipositor in
coriacea.

A relation between the variation described above and the geographic distribu-
tion seems unreliable. However, the variation is gradual and no gap of any
importance could be found. We are, of course, dealing with isolated populations
living in numerous islands, which might explain the observed variation.

Some previous records should be discussed. The one from Ceram appears to be
based on Walker’s record of lanceolata (Walker, 1870: 437). As the specimen is
now assigned to nubila, the occurrence of coriacea in Ceram needs to be
confirmed. De Haan’s record (1843: 214) of lanceolata from “Timor, Java” should
be considered unreliable as it was done by Karny (1931: 83). De Haan’s material
examined consists of 24 29 (RNH), all labelled “Java”. One pair agrees with
coriacea (as from Ambon), the other pair with nubila (as from the Moluccas). The
occurrence of coriacea in Celebes should be confirmed, as it was based on 29 of
doubtful origin before me and another © recorded by Karny (1924: 151). The same
can be said of its occurrence in New Guinea, which is based on 14 39 from
Hollandia (Karny, 1924: 151) and 14 at hand of doubtful origin, labelled “Nieuw
Guinea”. All Hollandia and other material from New Guinea before me belongs to
nubila.

Sexava nubila (Stäl, 1874)
(pl. 1 figs. 5, 6, pl. 2 figs. 7—9, pl. 3 figs. 15, 16, 20, 21, pl. 4 figs. 26, 27, map 1)

Moristus nubilus Stal, 1874: 96.

Sexava lanceolata (nec Stoll); Walker, 1870: 437.
Locusta lanceolata; De Haan, 1843: 214 (partim).
Sexava nubila; Kirby, 1906: 359.

Material studied: 9 holotype, labelled: 292 76 (NR) (parts of antennae lacking,
apex of fore wings slightly damaged).

Additional material: Celebes, Manado, i.1932, A. Reyne (14 BMNH); Talaud
Is., A. Reyne, ex coconut palms (14 19 BMNH), iv.1949, C. Franssen, on
coconut, ex coll. S. Leefmans (13 19 ITZ), Beo, 14—21.vi.1930, Snellius Exp. (6g
39 RNH); Manipa, 1864, Hoedt (1 RNH); Ceram (19 BMNH), Rukuwa, 30 km
E. of Amakar Kowsuto, 3.v.1970, R. F. Ellen (14 19 BMNH); Misool, 1870 (19
ITZ); Kei Is., C. Ribbe, coll. Br. v. W. (14 NMW), Kühn, coll. Br. v. W. (19

223

WILLEMSE: Sexavae of Melanesian Subregion

‘Sa19ads papxag Jo UONNINSIT

"| dew

iAuie} m

9981109 è
ejlqnu w
enexag

224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

NMW), 1903 (14 BMNH); Aru-Inseln, C. Ribbe, coll. Br. v. W. (13 19 NMW);
Banda Is., 5° S, 130° E, 6.iv.1975, J. E. Lloyd (14 CW); West New Guinea:
Sekroé, iii.1897, Schaedler (33 RNH); Fak Fak, viii.1874, C. J. L. Palmer (44 39
RNH); Poel Pandjang or Noha, viii.1904, Jhr. v. Nanhuijs (19 RNH); Sorong,
1.11.1929 (14 29 CW) & entre Sakoemi et Moemi, 13.11.1929 (19 CW) & Manoi
— Salawati, 2.iii.1929 (24 CW), Prince Léopold; Sedorfojo, vii.1952 (13 RNH) &
Seribaai, 5.viii.1952 (14 RNH), Mevr. Marcus v. d. Nieuwenhuizen; Komara,
ii.1963, Hr. & Mevr. Marcus v. d. Nieuwenhuizen (14 RNH); Jef Lie, 18.11.1957
(32 29 CW) & Sansapor, vi.1961 (13 19 CW) & Mega, 22.vii.1961 (1g CW) &
Koor, 26.vii.1961 (14 CW), R. T. Simon Thomas; Andai, acq. 1870, Rosenberg
(19 RNH); Paniai, 27.x1.1939, Nieuw Guinea Exp. N.A.G. 1939 (13 RNH);
Bernhard camp, 50 m, vii.1938, J. Olthof, Neth. Ind. Amer. New Guinea Exp. (ld
19 RNH); Maffin Bay, vi.1944, E. S. Ross (1 39 CAS); Hollandia, 1955—1956,
A. Klaassen (19 RNH), Ned. Nw. Guinea Exp. 1911, Dr. P. N. v. Kampen (34 69
RNH), vii.1938, L. J. Toxopeus, Neth. Ind. Amer. New Guinea Exp. (14 RNH);
Zoutbron, vi—vii.1911, Ned. Nw Guinea Exp. 1911, Dr. P. N. v. Kampen (1 9
RNH); Tanah Merah baai (South New Guinea), 19.viii.1910, Ned. Nw Guinea
Exp. 1911, Dr. P. N. v. Kampen (lg RNH); Mindiptana, 26.xi.1958, Br.
Monulphus (1 ¢ CW, 19 RNH); Mariang a. d. Digoel, 10 m, 12.ix.1959 (14 RNH)
& Kouh a. d. Digoel, 8.ix.1959, 10 m (84 79 RNH) & Kowage, 10 m, 8.ix.1959
(63 19 RNH) & Hijob, 25 m, 10.ix.1959 (13 29 RNH), Neth. New Guinea Exp.
Star Range; Java (De Haan vidit) (14 19 RNH); Sepik District (East New
Guinea), Bainyik, 1 .xii.1959 & 11.xii.1953, J. H. Ardley (29 DASF).

Stal described the species after a female. Sjostedt (1933: 14, pl. 25 fig. 1),
however, figured a male as Stäl’s type. Dr. T. Kronestedt (in litt. 24.111.1976)
informed me as follows:

“1. The specimen which Sjöstedt figured is the only one placed under the label
Sexava nubila in our collection, and it is obviously the same specimen as he
figured.

2. The same drawer contains also 3 specimens placed under coriacea L., two of
which bear a name label “coriacea”, one without any label. ... Assuming that the
one lacking any label is the type of nubila, we compared the measurements given
in Stäl’s original description with the measurements given in his redescription of
coriaceus. It then turned out that Stäl’s measurements given for coriaceus fitted the
specimen which he had labelled as such and his measurements given for nubila
fitted the specimen without labels.

3. The male specimen labelled nubilus bears labels in Stäl’s handwriting ‘“Moristus
Stal” and “nubilus Stal” as well as a label ’’Platyphyllum coriaceum (?) L. Serv.”
and a red printed label “Typus”, the latter apparently put on by Sjostedt.”

Presumably, when Sjostedt figured the type of Stäl’s nubilus, the male
erroneously beared Stäl’s labels instead the female. The latter specimen is before
me and agrees fully with Stäl’s description. It is considered the holotype of
Moristus nubilus (pl. 2 fig. 7).

The additional material at hand from New Guinea, Kei and Aru Is. and the
southeastern Moluccas agrees with the holotype. The particular form of the

WILLEMSE: Sexavae of Melanesian Subregion 225

species occurring in Talaud and Nanusa Is. is discussed below.

The species is defined as indicated in the key. Quite characteristic are the shape
of the male subgenital plate (pl. 3 figs. 15, 16), the cercus (pl. 3 figs. 20, 21) and the
stridulatory file (pl. 4 figs. 26, 27). The latter, compared with coriacea, is less
fusiform and arcuate, 5.5.-7.0 mm long, number of teeth 54-65 of which the
anterior 10-20 are fine and about blunt, covering the anterior sixth to eighth part
of the file. Width of the file strongly increasing in the anterior part, reaching
maximum 0.6-0.8 mm in the middle third, decreasing posteriorly but slightly.
Spacing of the teeth wider than in coriacea, moderately increasing posteriorly,
distance between successive posterior teeth 1-2 times as large as between the
anierior sharp teeth. Mirror and its frame of the right male fore wing slightly
different from that in coriacea, but the available material of the latter does not
allow reliable conclusions.

Variation. In spite of individual variation, the shapes of the male subgenital plate,
cercus and stridulatory file, are rather uniform throughout the material at hand,
including that from Talaud Is. The same can be said of the female subgenital plate,
figured by Leefmans (1927b: pl. 5 fig. 3). Material from New Guinea (pl. 1 fig. 5,
pl. 2 fig. 8), Kei and Aru Is. and the southeastern Moluccas is uniform in general
appearance (slender) and coloration (brown). However, that from Talaud Is. (pl. 1
fig. 6, pl. 2 fig. 9), Nanusa Is. (cf. Leefmans, 1927b and the male from northern
Celebes differ in more robust general appearance, wider thorax, wider and
comparatively shorter wings and often green general colour. The lower sides of the
femora of a male from Banda are dark brown, but not solid black. In all material
at hand, usually a series of distinct dark brown flecks with a central yellow dot on
the radial area of the fore wing is present.

Measurements: body ¢ 55—72, 9 50—70; fore wing & 68—80, 9 75—86; hind
femur ¢ 35—43, 2 38—43; ovipositor 30—43.

Distribution (map 1). The range covers northeastern and all western New
Guinea, Kei and Aru Is., southeastern Moluccas, extending into the Talaud and
Nanusa Is. and (?) northern Celebes.

Localities: Celebes: Manado; Talaud and Nanusa Is. (Leefmans, 1927b: 13;
Oudemans, 1927: 267; Franssen, 1954: 99-102; Reyne, 1960: 232); Manipa
(between Buru and Ceram); Ceram (Walker, 1870: 437), East Central and Watai
(Karny, 1924: 151), Rukuwa; Banda; Misool; Kei Is. (Redtenbacher, 1892: 201;
Karny, 1926: 183); Aru Is. (Redtenbacher, 1892: 201); West New Guinea: Sekroë;
Pandjang I.; Sorong (C. Willemse, 1933: 9); Sakoemi-Moemi; Manoi-Salawati (C.
Willemse, 1933: 9); Sedorfojo; Seribaai; Komara; Jef Lie; Sansapor; Fak Fak;
Mega; Koor; Andai; Manokwari (C. Willemse, 1933: 9); Paniai; Bernhard Camp;
Maffin Bay; Hollandia (Karny, 1924: 151); Zoutbron; South New Guinea; Tanah
Merah baai; Sud-Neuguinea (Karny, 1924: 151); Frederik-Hendrik I. (Karny, 1924:
151); Mindiptana; Mariang; Kouh; Kowage; Hijob. East New Guinea: Sepik distr.,
Bainyik.

Discussion. The species is well-defined in the male. The female of the form

occurring in the Talaud and Nanusa Is., however, resembles strongly that of
coriacea. Their distinction is discussed above under the latter species. Besides, the

226 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

female of nubila resembles superficially that of Segestidea novaeguineae. The latter
differs in more oblique course of Cul of the fore wing, stronger ventral spines of
the hind femur and in colour pattern of the hind leg.

A record of nubila from Java (K. K. Hofmuseum Wien) by Redtenbacher (1892:
201) is unreliable (Karny, 1924: 151). His material could not be traced in the
Vienna museum (Kaltenbach, in litt. 12.xi.1975). The pair labelled “Java” of De
Haan is discussed under coriacea, and the locality is considered unreliable. Karny’s
record (1924: 151) from Batjan was based on 19 while 1g 29 of the same series
(originating from Leefmans) were assigned to coriacea. | doubt the correct
identification of the “nubila’ female. The occurrence of nubila in Batjan should be
confirmed, based on a male. Of zoogeographic interest is the Manado male (N.
Celebes), which agrees fully with the nubila form of the Talaud and Nanusa Is. This
single record needs confirmation as the locality appears isolated from the known
range of the species (map 1).

Sexava karnyi Leefmans, 1927
(pl. 2 figs. 10, 11, pl. 3 figs. 17, 22, pl. 4 fig. 28, map 1)
Sexava karnyi Leefmans, 1927a: 411, figs. 1-5 (type-locality: Poat I.).

Material studied: Kp. Baroe (Ampana M. Cel.), viii.1949, A. Cohen, on coconut
(1g); Posso, ix.1948, Dr. C. Franssen, on coconut leaf (19 ) (both ex coll. Dr. S.
Leefmans, ITZ).

The types of this species could not be traced. The material before me (pl. 2 figs.
10, 11) agrees completely with the description and figures of karnyi. The fore wing,
which is wide and rather tapering apically, shows the venation characteristic in
Sexava.

The species is defined in the key. The stridulatory file (pl. 4 fig. 28) of the
available male is 6.5 mm long, number of teeth 60, of which the anterior 35 about
blunt and covering the anterior fourth of the file length. Width of the file
increasing in anterior half, reaching a maximum of 0.4 mm in middle, decreasing
posteriorly to 0.2. mm. Spacing of the teeth conspicuously increasing posteriorly,
greatest distance between successive posterior teeth about 10 times more than
between the anterior sharp teeth. Mirror and its frame of right male fore wing
much as in the other two species of the genus. Flexed wings reach the apical third
of hind tibia and tip of ovipositor. The male subgenital plate (pl. 3 fig. 17;
Leefmans, 1927a: fig. 2) resembles much that in nubila. The male cercus (pl. 3 fig.
22) has a robust and short apex, bearing a minute spine on its anterior edge. The
female subgenital plate of the available specimen appears more widely emarginate
than in the figure by Leefmans (1927a: fig. 3). The coloration of the pair agrees
fully with the original description: general colour brown, lower and inner side of
proximal part of hind femur conspicuously solid black.

Measurements (partly after Leefmans): body g 53—58, 9 48—54; fore wing &
52—53, © 56—60; hind femur ¢ 36—40, © 38—39; ovipositor 26—27.

Distribution. Known only from Togian Is. and the southern opposite part of
Celebes (map 1).

WILLEMSE: Sexavae of Melanesian Subregion 227
Localities: Poat I. (Leefmans, 1927a: 412); Celebes: Posso; Ampana.

Discussion. The species is well-defined, especially by the male stridulatory file
and the coloration of the hind femora. Variation is insufficiently known.
Previously recorded only from the type-locality.

Segestes Stal, 1877

Segestes Stal, 1877: 45; Redtenbacher, 1892: 189, 197.
Type-species by monotypy: Segestes vittaticeps Stal, 1877.

Segestes is characterized, as noted in the key, by lacking the dorso-apical spines
of fore and mid tibiae in combination with usual course of Cul of the fore wing,
delimiting a well-defined triangular area which, in the male, contains the main
attributes of the stridulatory apparatus. As pointed out above under the generic
characters, exceptionally a dorso-apical spine has been found on the fore and mid
tibiae of some specimens considered Segestes decoratus.

Up to now, the following taxa have been arranged under Segestes:
vittaticeps Stal, 1877
punctipes Redtenbacher, 1892
unicolor Redtenbacher, 1892
fuscus Redtenbacher, 1892
decoratus Redtenbacher, 1892
frater Hebard, 1922
sp. Leefmans, 1927
celebensis Karny, 1931
beieri Kastner, 1934
acuminatus Kastner, 1934
grandis C. Willemse, 1955 (correctly assigned to Sexava by C. Willemse, 1957)

The latter taxon is synonymized with Sexava coriacea (see above) and acuminatus
is now transferred to Segestidea. Of the other species three occur in the
Philippines, one in Palau I., one in Celebes, one in Obi and one in New Guinea.
The typespecies, unicolor from Palau I. and decoratus from New Guinea, will be
discussed, while the other species are shortly memorized only. Three new species
from New Guinea will also be described. Provisional keys to previously recognized
species were given by Redtenbacher (1892: 198), Kastner (1934: 46), and C.
Willemse (1961: 107). When using these keys, it is noted here that styli of the male
subgenital plate of vittaticeps were incorrectly thought to be lacking. A key to the
species occurring in New Guinea is given below.

Segestes vittaticeps Stal, 1877
(pl. 6 fig. 30, pl. 7 fig. 34, pl. 9 figs. 48, 53, pl. 10 fig. 64)
Segestes vittaticeps Stal, 1877: 45.

The species is known from the syntypes (14 19 RN), 1g (NMW) recorded by

228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

Redtenbacher (1892: 198) (misidentification?) and 14 29 (ANSP) recorded by
Hebard (1922: 176). The syntypes were figured by Sjostedt (1933: 14, pl. 22 figs. 2,
3) and the male subgenital plate of Redtenbacher’s male by Kastner (1934: 47, fig.
14).

The male syntype is before me and hereby designated lectotype (pl. 6 fig. 30,
pl. 7 fig. 34). It is labelled: Ins. Philipp., Semper, Segestes vittaticeps Stàl, Typus.
The specimen agrees with Stäl’s description and Sjöstedt’s figure, but lacks part of
the stridulatory file, which was not so in Sjostedt’s figure, and the position of the ~
legs differs also slightly from that figure.

Redescription.

Small (pl. 6 fig. 30, pl. 7 fig. 34). Fastigium of vertex obtusely pointed, reaching
about middle of scape, shallowly sulcate. Pronotum with dorsum somewhat
flattened medially, slightly rounded laterally, anterior margin slightly convex,
posterior margin straight; lateral lobe slightly longer than high, deepest point of
lower margin about in the middle, from there rounded, posteriorly more so than
anteriorly.

Flexed wings extending just behind hind knee. Fore wing narrow, in distal half
gradually tapering toward narrowly rounded apex; archedictyon well developed,
membrane more or less opaque; Cul running obliquely toward hind margin of
wing, almost reaching the latter at short distance from wing-base and from there
parallel and close to hind margin of wing, vanishing into archedictyon about
middle of wing length; stridulatory area well-defined, file partly lacking; mirror
(pl. 10 fig. 64) twice as long as wide, elongate-elliptical, fold extending slightly over
mirror, its outline almost straight and parallel to anterior margin of mirror.

Fore and mid femora unarmed. All knee-lobes with one spine. Fore tibia
without dorsal spines, mid tibia with 2 dorsal spines on distal half of posterior
margin, no dorso-apical spines. Apical half of hind femur with series of ventral
spines.

Cercus (pl. 9 fig. 53) almost rectangularly incurved in apical fourth, gradually
tapering to a short tooth. Subgenital plate (pl. 9 fig. 48) strongly elongate, slightly
narrowing apically, apex divided by comparatively deep, narrowly parabolic
incision into pair of lobes, tips of the latter truncate with distinct styli.

General colour pale brown (‘‘olivaceo-virescens’’ in Stäl’s description). Median
black stripe over occiput, narrowing anteriorly and reaching fastigium of vertex.
Hind margin of fore wing narrowly yellowish-white. Lower and inner sides of
proximal half of hind femur solid black. Spines of legs with tips black, of hind
femur black at their bases.

Measurements: body 39; fore wing 33; hind femur 28.

Distribution. As far as now only known from the Philippine Is.: — (Stäl, 1877:
45; Redtenbacher, 1892: 198); Surigao, Mindanao (Hebard, 1922: 176).

Discussion. Redtenbacher and Kästner based their diagnosis of vittaticeps on a
male in the Vienna museum. The subgenital plate of this specimen, figured by
Kastner, lacks the styli, a character used in their keys to the species. Whether this
difference with Stäl’s type is of much importance is an open question.

WILLEMSE: Sexavae of Melanesian Subregion 229

Segestes punctipes Redtenbacher, 1892

Segestes punctipes Redtenbacher, 1892: 199.

Known only from the © holotype (NMW). The species is discussed by Hebard
(49223170):

Distribution. Philippine Is., without precise locality.

Segestes fuscus Redtenbacher, 1892
Segestes fuscus Redtenbacher, 1892: 199.

Known only from the 9 holotype (NMW).
Distribution. Philippine Is., without precise locality.

Segestes celebensis Karny, 1931

Segestes celebensis Karny, 1931: 72, fig. 37.

Know only from the ¢ holotype and | juvenile 9 (depository?).
Distribution. Celebes: Matinang-Kette & Minahassa, Karowa-Popo.

Segestes beieri Kastner, 1934

Segestes beieri Kastner, 1934: 48, figs. 15—17.

Known only from the syntypes (1¢ 19 NS).
Distribution. Philippine Is.: Mindoro (Kastner, 1934).

Segestes frater Hebard, 1922
Segestes frater Hebard, 1922: 177, pl. 16 fig. 2.

Known only from the © holotype (ANSP) and 14 (NS). The latter was
described by Kastner (1934: 51, figs. 20, 21). I have before me a female from
Laiwui, Obi, 29.ix.1953 (CW), which agrees with Hebard’s description except for
the coloration, which is unicolorous green.

Distribution. Known only from the Moluccas: Obi (Hebard, 1922: 177; Kastner,
1934: 51).

Segestes sp. Leefmans, 1927

Segestes sp. Leefmans, 1927b: 15, pl. 5 fig. 4, pl. 6 fig. 4.

According to Leefmans (1927b) and Karny’s opinion, a not yet described species
from Loleba I., near Halmaheira. I could trace only one discoloured female in bad
condition, labelled: Segestes spec. Loleba I. (Halmaheira) sent by Leefmans
(BNNH). The specimen agrees with Leefmans’ data but by lack of the male,
further comments are postponed.

230 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

Segestes unicolor Redtenbacher, 1892

(pl. 6 fig. 32, pl. 7 fig. 35, pl. 9 fig. 54, pl. 10 fig. 59)
Segestes unicolor Redtenbacher, 1892: 199.

Known from the 9 holotype (NMW) and another 9 (BPBM) recorded by
C. Willemse (1951: 342). I have before me 2¢ labelled: West Caroline Is., Koror,
Palau, 1.xii. & 20.x.1971, M. R. Lundgren (CAS). Presumably, on the basis of
topographic evidence, these males represent unicolor.

Description of male.

Small (pl. 6 fig. 32, pl. 7 fig. 35). Head, thorax and wings as in vittaticeps.
Stridulatory file (pl. 10 fig. 59) slightly arcuate and fusiform, 2.9—3.0 mm long,
number of teeth 100—110, of which anterior 20—30 ones fine, blunt and covering
anterior sixth of file length. Width of file reaching maximum 0.2 mm at the end of
proximal fourth of file length, decreasing slightly posteriorly. Spacing of teeth
almost regular. Stridulatory area of right fore wing as in vittaticeps.

Anteroventral margin of fore and mid femora with 1—3 spines in apical half.
Knee-lobes with one spine. Fore tibia without dorsal spines, posterodorsal margin
of mid tibia with 5 spines, no dorsoapical spine. Hind femur with series of ventral
spines.

Cercus (pl. 9 fig. 54) slender, incurved in apical fourth, rather abruptly tapering
to short, slightly incurved apical tooth. Subgenital plate as in vittaticeps.

Unicolorous pale brown or pale green, except for yellowish-white hind margin
of fore wing and spines of legs, which are completely black or with tips black.

Measurements: body 36—37; fore wing 38—39; hind femur 27—28.

Distribution. Known only from Pelew (Redtenbacher, 1892) or Palau I. (C.
Willemse, 1951), West Caroline Is.

Discussion. As far as can be judged now, the species differs from the type-
species mainly in the shape of the male cercus and the uniform coloration. A
record under the name unicolor by Karny (1924: 150) from Neu-Pommern (= New
Britain) probably does not refer to this species, but to Segestes decoratus.

Among the material before me from New Guinea and the Bismarck Archi-
pelago, four species of Segestes can be recognized: decoratus Redtenbacher,
cornelii sp.n., stibicki sp.n., and brevipennis sp.n.

Key to the species of Segestes from New Guinea and the Bismarck Archipelago
1. Fastigium of vertex (pl. 9 figs. 46, 47) long, reaching at least apical margin of
scape, apex acute, thorn-like; apex of fore wing wide, more or less truncately
rounded; knee-lobes usually with one spine; ¢ subgenital plate with or without
Styl RE ERENTO DDR ID IST SIE) A 2
— Fastigium of vertex (pl. 9 fig. 45) shorter, by far not reaching apical margin of
scape, apex obtusely pointed; apex of fore wing narrowly rounded; knee-lobes
usually with two spines; ¢ subgenital plate with styli, cercus, stridulatory file

WILLEMSE: Sexavae of Melanesian Subregion 231

and mirror with frame as in pl. 9 figs. 49, 55, pl. 10 figs. 60, 65 (NW and E. New
Guinea, western New Britain) .............. decoratus Redtenbacher
2. Larger, wing length g © 57—64 mm (pl. 6 fig. 31, pl. 7 fig. 40); & subgenital
plate without distinct styli, cercus, stridulatory file and mirror with frame as in

pl. 9 figs. 50, 56, pl. 10 figs. 61, 66 (East New Guinea) ....... cornelii sp.n.
— Smaller, wing-length & 9 32-47 mm (pl. 7 figs. 38, 39, 41, pl. 8 figs. 42—44);
desubgenitaliplateswathsstylikasbingpl Sri g ss 2 iene renee ts ce a 3

3. Fastigium of vertex reaching 2nd—3rd antennal segment (pl. 9 fig. 47); g
subgenital plate with narrow, parallel-sided apical incision and distinct styli (pl.
9 fig. 51); g stridulatory file about 2.4 mm long with about 150 teeth (pl. 10 fig.
62) (East New Guinea: Madang District) an anne. stibicki sp.n.
— Fastigium of vertex shorter, reaching apical margin of scape; & subgenital
plate with V-shaped apical incision and minute styli (pl. 9 fig. 52);
3 stridulatory file about 2 mm long with about 71 teeth (pl. 10 fig. 63) (West
ING WEG WINE A) PEU ra DAR GRIME ele Ce tc ONG Me brevipennis sp.n.

Segestes decoratus Redtenbacher, 1892
(pl. 6 fig. 33, pl. 7 figs. 36, 37, pl. 9 figs. 45, 49, 55, pl. 10 figs. 60, 65, map 2)

Segestes decoratus Redtenbacher, 1892: 18, fig. 4; Kastner, 1934: 52.

Segestes unicolor; Karny, 1924: 150(?).

Eumossula gracilis; C. Willemse, 1958: 122 (only Bubia); F. Willemse, 1966: 48 (same).
Segestidea insulana; F. Willemse, 1966; 48 (only Bubia).

Sexava femorata; F. Willemse, 1966: 49 (only 19 Manam ].).

Sexava species B: O'Connor, 1959: 122.

This species was badly known. It was described after a pair from ‘‘Neu-Guinea”’.
The male could not be traced in the Vienna museum (Kaltenbach, in litt.
11.11.1976). The female is before me and is considered holotype (pl. 7 fig. 36).
Kästner (1934) recorded additional material from ‘‘Neu-Guinea’’, but gave no
further comments. The material listed below agrees with the holotype.

Material studied: 9 holotype, labelled: Coll. Br. v. W. Neu-Guinea Fruhstorfer,
det. Br. v. W. Segestes decoratus Redt., Typus, 18 366 (NM W) (lacks parts of tarsi;
left side of head and thorax slightly crushed).

Additional material: N. Nov. Guinea, Boven Sermowai, NNW 400 m, 1—
8.vi.1911, N.K.G. (= NW New Guinea) (13 RNH); Gulf District, Murua Agric.
Stat. near Kerema, vii.1959, F. X. Rayan, on cocos (14 19 DASF); Northern
District, Popondetta, Casey’s Plantation, xi.1960 & 10.v.1960, G. S. Dunn, under
the bark of bush trees (14 19 DASF); Morobe District, Siassi I., 6.xii.1969, A.
Hinton, defoliating coconut palms (14 DASF); Agric. Exp. Stat. Bubia near Lae,
28.11.1955, J. Szent-Ivany, on coconut frond (14 Segestidea insulana det. C.
Willemse 1958 & 13 19 Eumossula gracilis det. C. Willemse 1958, CW); Bubia
Agric. Stat., 20.11.1957, Luna de Carvalho (19 BMNH); Bubia via Lae,
12.vii.1960, A. Catley, feeding on Cocos nucifera (54 59 CW); Deutsch Neu-
Guinea, Sattelberg, H. Rolle, Berlin (1 ¢ CW); Siki via Finschhafen, 20.vii.1960,
A. Catley, ex Cocos nucifera (1g DASF); Madang District, Manam I., Baliau

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

232

‘ds «

19301) +

sıuuadınaıq %

I19U109 w

ıyDıqııs B

sn}e1099p è
sajsabas

'sar9ads sajsadag Jo vornginsig ‘7 dep

WILLEMSE: Sexavae of Melanesian Subregion 233

Village, 26.ix.1960, J. I. Cox (19 Sexava femorata det. C. Willemse, CW); Awar
Plantation, via Bogia, xi.1960, J. I. Cox (34 29 CW); Madang, 4.v.1932, J. L. F.,
coconuts {14 BMNH); West New Britain, Bitokara C. M., Talasea, 26.11.1971, T.
Laklo, on coconuts (59 DASF); Talasea, Volupai Plantation, 10.v.1966. D. F.
O’Sullivan (19 DASF); West New Britain, Lingalinga Plantation, vi.1959, J. H.
Barrett, on Theobroma (19 DASF); West New Britain, Wakanaka Village,
23.1.1974, C. H. Perry, resting in fronds of coconut (1 9 DASF).

Redescription.

d (pl. 6 fig. 33), large. Fastigium of vertex (pl. 9 fig. 45) extending but slightly
beyond antennal scrobae, reaching middle of scape or shorter, apex obtusely
pointed, often with shallow furrow. Pronotal dorsum strongly rounded laterally,
anterior margin slightly convex, posterior one straight. Lower margin of pronotal
lateral lobe obtuse-angulately rounded, deepest point just distally of middle of
lobe length, from there slanting upwards, posteriorly steeper than anteriorly.

Flexed wings reaching middle of hind tibia or slightly shorter. Fore wing narrow,
tapering gradually towards narrowly rounded apex; moderate archedictyon.
Stridulatory file (pl. 10 fig. 60) scarcely arcuate, 3.5—4.0 mm long, number of teeth
183—196, of which anterior 50—60 fine, more or less blunt and covering anterior
fifth to sixth of file length, the other 130—140 ones sharp. Width of file reaching
maximum 0.4—0.5 mm at end of proximal fourth of file length, decreasing
posteriorly to one-third of maximum width. Spacing of teeth slightly increasing
posteriorly, comparatively more in anterior than in posterior part of file. Mirror
(pl. 10 fig. 65) about twice as long as wide, roughly elliptical with antero-apical
margin angulate; fold moderately extending over mirror, outline sinuate.

Fore femur with 0—3, mid femur with O—1 spines on anteroventral margin. All
knee-lobes with a larger dorsal and a smaller ventral spine; occasionally hind
knee-lobe with three and fore knee-lobe with one spine. Fore tibia with O—1, mid
tibia with 5—8 dorsal spines on posterior margin; dorso-apical spines usually
lacking.

Cercus (pl. 9 fig. 55) incurved, tapering apically to short and distinctly incurved
tooth. Subgenital plate (pl. 9 fig. 49) almost four imes as long as smallest width,
lateral margin slightly concave in the middle; apex divided by arrowly parabolic
incision into pair of lobes with obliquely truncate or slightly sinuate tips. Stylus
comparatively long, inserted in lateral half of tip of lobe and usually pointing
inward.

General colour green or, less often, brown. Antennae unicolorous or very
slightly annulated pale and dark. Palpi from orange via yellowish to pale brown.
Head and pronotum unicolorous, but usually with pale orange-yellow stripe
running from behind eye toward antero-ventral angle of pronotum, extending over
lower margin of pronotal lateral lobe and similarly coloured median stripe from
occiput over pronotal dorsum. Lateral stripes may extend over pleurae and
median one over cubito-anal areas of fore wing. Fore wings of general colour,
immaculate, membrane ranging from little transparent to slightly opaque. Legs of
general colour or yellowish, lower and partly inner side of hind femur orange in

234 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

green specimens, reddish brown in brown specimens. Knees either of general
colour or black from below, or sometimes completely black. Spines of legs with
tips black or black basally. Tibiae and tarsi of general colour, tarsi dull brown from
below or sometimes completely dark brown.

Q (pl. 7 figs. 36, 37), larger than male. Wings comparativels long, as in male.
Ovipositor with upper margin straight, reaching middle of hind tibia, in dorsal view
covered by flexed wings or almost so. Subgenital plate wide, triangular, with
median apical emargination. Coloration as in male, basal part of hind margin of
fore wing often narrowly orange or yellow.

Variation. It is of particular interest to note that in some specimens the left or
right fore and mid tibiae have a posterior dorso-apical spine. This was found in
three out of seven females, all from New Britain. Otherwise the specimens agree
fully with the material from New Guinea. Whether the instability of this generic
character is limited to the population of New Britain is an open question. It is
advisable to study the associated males from New Britain to establish the
conspecificity with decoratus.

Measurements: body & 51—54, 9 56—61; fore wing & 52—59, 9 63—72; hind
femur & 32—37, Q 39—42; ovipositor 32-35.

Distribution. The range covers the eastern half of the mainland of New Guinea,
extending into some islands near its coast and New Britain (map 2).

Localities: Neu-Guinea (Redtenbacher, 1892; Kastner, 1934); West New
Guinea: Sermowai near Hollandia; East New Guinea, East Sepik District: Mushu
I. (O'Connor, 1959); Madang District: Madang; Awar Plantation near Bogia
(O’Connor, 1959); Baliau Village, Manam I.; Gulf District: Murua Agric. Stat.
near Kerema; Northern District: Casey’s Plantation near Popondetta; Morobe
District: Siki near Finschhafen; Bubia near Lae and Agric. Stat. Bubia; Sattelberg;
Siassi I.; West New Britain District: Bitokara C. M. near Talasea; Volupai
Plantation near Talasea; Wakanaka Village; Lingalinga Plantation.

Discussion. The species is well-defined and easily distinguished from other
Segestes species. However, decoratus may resemble some species of Segestidea,
especially uniformis (C. Willemse) and gracilis (C. Willemse). As noted above, in
some specimens of decoratus from New Britain dorso-apical spines of fore and mid
tibiae are present, which is misleading as to correct generic assignment. In these
cases, the following notes may be helpful.

Segestidea uniformis (C. Willemse), which occurs in the Admiralty Is. (Manus
District), has the pronotal lateral lobe narrower, its lower margin not yellow but of
similar colour as the lobe and the deepest point not beyond but about midway the
length. The male subgenital plate (pl. 15 fig. 93) is narrower and the male cercus
(pl. 15 fig. 101) shorter. The male stridulatory file is about similar but the number
of teeth is smaller (130—140 versus 183—196) and the spacing of he teeth more
widely (pl. 16 fig. 109); the mirror is distinctly less covered by the fold (pl. 18 fig.
117). Head, thorax, wings and legs are unicolorous.

Segestidea gracilis (C. Willemse), both in the nominate subspecies (known from
New Ireland District) and in simulatrix ssp.n. (known from New Britain), has the
wings shorter and the ovipositor longer (pl. 12 figs. 77—79, pl. 13 figs. 80—82). The

WILLEMSE: Sexavae of Melanesian Subregion 235

lower margin of the pronotal lateral lobe is not yellow but of general colour,
slightly convex and not at all obtuse-angulately rounded. The male subgenitel
plate and cercus are smaller (pl. 15 figs. 94, 95, 102, 103). The male stridulatory file
in simulatrix is of similar shape, but the number of teeth is much smaller (117—139
versus 183-196) and their spacing much wider (pl. 16 fig. 111), while the shape of
the file in nominate gracilis is quite distinct, its anterior half being much narrower
(pl. 16 fig. 110). Head, thorax, legs and wings are unicolorous.

Karny (1931: 72), who had the species not before him, doubted as to the correct
arrangement of decoratus under Segestes, based on the number of spines of the
knee-lobes and the presence of styli of the male subgenital plate. The material at
hand demonstrates clearly the variability of the number of spines, and the styli are
present in the type-species of Segestes. Karny’s presumption may be, however, of
some importance, but for other reasons, viz., the occasional presence of the fore
and mid tibial dorso-apical spines. Material recorded under Sexava species B by
O’Connor (1959) was not available, but certainly refers to decoratus. Karny’s
record of Segestes unicolor Redtenbacher from Neu-Pommern (= New Britain)
may refer to decoratus rather than unicolor, which is known only from the West
Caroline Island Palau. I have not examined his material.

Segestes stibicki spec. nov.
(pl. 7 figs. 38, 41, pl. 8 figs. 42, 43, pl. 9 figs. 47, 51, 57, pl. 10 figs. 62, 67, map 2)

The species is named after Dr. J. L. Stibick, whose activities urged me to this
study.

Material studied: ¢ holo-, 8 © paratypes, labelled: Madang District, Kaironk
area, xii.1971-1.1972, J. Menzies (¢ holo-, 6 9 paratypes CW; 2 © paratypes
DASF) (depositories proposed by Dr. J. Stibick) (holotype lacks both antennae,
both fore tibiae, left hind leg, right hind tibia and apex of left fore wing).

Description.

& (pl. 7 fig. 38, pl. 8 fig. 42), small. Face rather reclinate. Fastigium of vertex
(pl. 9 fig. 47) thorn-like, apex acute, extending far beyond antennal scrobae,
reaching third antennal segment (first segment of flagellum), in profile slightly
upcurved. Pronotum short; dorsum flattened, weakly rounded laterally, ‘‘shoul-
ders’ distinct, anterior margin almost and posterior margin quite straight, both
with weak median tubercle; lateral lobe about as long as high, lower margin
slightly rounded, deepest point at or just before middle of length.

Flexed wings reaching distal end of proximal third of hind tibia. Fore wing
narrow, margins about parallel, slightly tapering toward obliquely truncate apex.
Stridulatory file (pl. 10 fig. 62) slightly arcuate and fusiform, 2.4 mm long, number
of teeth 150, of which anterior 40 fine and covering anterior seventh of file length,
other 110 sharp. Width of file reaching maximum 0.26 mm at end of anterior
fourth, decreasing posteriorly to about half maximum width. Spacing of teeth
slightly increasing posteriorly, comparatively more in anterior part. Mirror (pl. 10

236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

fig. 67) about twice as long as wide, roughly elliptical, fold extending well over
mirror, outline convex.

Fore and mid femora with 0—5 spines on anteroventral margin. All knee-lobes
with one spine. Fore and mid tibiae with 2-5 dorsal spines on the anterior and 0-3
dorsal spines on posterior margin, no dorso-apical spines.

Cercus (pl. 9 fig. 57) slender, slightly incurved, tapering to a short tooth.
Subgenital plate (pl. 9 fig. 51) very narrow, seven times as long as smallest width,
apex divided by narrow, about parallel-sided incision into pair of narrow lobes
with rounded tips, each with short stylus.

© (pl. 7 fig. 41, pl. 8 fig. 43). Apex of flexed wings reaching tip of ovipositor
or almost so. The ovipositor slightly upcurved, comparatively short, reaching not
quite middle of hind tibia. Subgenital plate wider than long, with slight median
apical incision, lobes angulately rounded. Lower margin of ninth tergite extending
over upper part of gonangulum (= valvifer), forming shallow groove.

General colour pale brown or pale green. Antennae more or less darkly
annulated. Apex and dorsal side of fastigium of vertex, four longitudinal lines over
occiput and lateral margins of pronotal dorsum dark brown; sometimes whole
occiput, pronotal dorsum and cubito-anal areas of fore wing dark brown. Fore
wing with more or less numerous scattered black dots, membrane partly
transparent and slightly opaque. Pleurae and coxae with or without some black
dots. Lower side of fore femur usually dark brown, fore tibia and mid leg of
general colour. Hind femur between bases of knee-lobes, often inner lower margin
and sometimes whole lower side, blackish brown. Hind tibia of general colour,
lower side often dark brown. Spines brown, tips black, those of hind femur usually
completely black.

Measurements: body ¢ 35, 9 36—38; fore wing ¢ 36, 9 36—41; hind femur G 22
Q 24—26; ovipositor 18—20.

Distribution. Known only from the type-series, East New Guinea, Madang
District: Kaironk area (map 2). .

Discussion. The species is well-defined, although at a first glance it resembles
Segestidea acuminata (Kastner). The latter species, known in the female sex only,
differs in presence of fore and mid tibial dorso-apical spines, larger measurements,
shorter fastigium of vertex and wider lobes of the female subgenital plate

No previous records.

Segestes brevipennis spec. nov.

(pl. 7 fig. 39, pl. 8 fig. 44, pl. 9 figs. 52, 58, pl. 10 fig. 63, map 2)

Material studied: & holotype, labelled: Egemendora, J. Eyma leg., Nieuw
Guinea Exp. K.N.A.G. 1939, mid Oct. 1939 (CW) (left mid leg lacking).

Description.
d (pl. 8 fig. 44), small. Fastigium of vertex thorn-like, extending well beyond
antennal scrobae, just reaching apical margin of scape, slightly upcurved. Pronotal

WILLEMSE: Sexavae of Melanesian Subregion 237

dorsum flattened in the middle, slightly rounded laterally, “shoulder” well
developed; anterior margin slightly convex, posterior margin straight, both
margins with weak median tubercle. Pronotal lateral lobe as long as high, lower
margin scarcely convex, angles rounded, deepest point about in the middle.

Flexed wings reaching distal end of proximal fourth of hind tibia. Fore wing
with margins roughly parallel, scarcely tapering toward obliquely truncated apex;
archedictyon well-developed. Stridulatory file slightly arcuate and fusiform (pl. 10
fig. 63), 2.0 mm long, number of teeth 71, of which anterior 17 fine, covering
anterior seventh of file length, the other 54 sharp. Width of file strongly increasing
in anterior seventh of file length, reaching there its maximum (0.35 mm),
decreasing posteriorly to about half maximum width. Spacing of teeth increasing
in anterior part. Mirror damaged (not figured), about as in stibicki.

Fore and mid femora with 2-3 spines on anteroventral margin. Fore tibia with 3
dorsal spines on anterior margin, posterior margin unarmed. Mid tibia with 2
dorsal spines on anterior and 6 dorsal spines on posterior margin. Fore and mid
tibial dorso-apical spines lacking. All knee-lobes with one spine.

Cercus (pl. 9 fig. 58) slightly incurved, tapering apically to a short tooth.
Subgenital plate (pl. 9 fig. 52) about six times as long as smallest width, apex
divided by wide V-shaped incision into pair of lobes with truncate tips, in the
middle of the latter a minute stylus.

General colour dark brown. Antennae slightly annulated pale and dark brown.
Fore wing with cubito-anal areas and several scattered points and dots dark
brown, hind margin narrowly yellowish brown. Lower side of all legs dark brown,
especially of hind femur and hind knee. Spines with tips black.

©. Unknown.

Measurements: body 34; fore wing 32; hind femur 21.

Distribution. Known only after the type, West New Guinea: Egemendora (map
DI

Discussion. The species comes near stibicki from which it is separated as
indicated in the key. Further material is needed to establish the reliability of the
differences mentioned.

No previous records.

Segestes cornelii spec. nov.
(pl. 6 fig. 31, pl. 7 fig. 40, pl. 9 figs. 46, 50, 56, pl. 10 figs. 61, 66, map 2)

The species is named after my late father, Cornelis J. M. Willemse.

Material studied: ¢ holo-, 34 49 paratypes, labelled: Kandep, West. Highl.
Distr., New Guinea, 14.11.1964, J. J. H. Szent-Ivany, on Sach. robosium and
Pandanus (holo-, 13 29 paratypes CW; 2g 29 paratypes DASF) (holotype lacks
tips of antennae only).

Description.
3 (pl. 6 fig. 31), large. Fastigium of vertex (pl. 9 fig. 46) thorn-like, apex acute,

238 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

reaching just beyond apical margin of scape, slightly upcurved. Pronotal dorsum
slightly flattened in middle, slightly rounded laterally, anterior margin almost,
posterior margin quite straight. Pronotal lateral lobe slightly longer than high,
lower margin slightly convex, deepest point about in middle.

Flexed wings reaching almost middle, or usually distal end of proximal third of
hind tibia. Fore wing wide, margins about parallel, slightly tapering toward a wide,
obliquely truncated and rounded apex; archedictyon well-developed, transverse
veins poorly developed. Stridulatory file (pl. 10 fig. 61) almost straight, 3.1—3.2
mm long, number of teeth 85— 100, of which anterior 25—35 very fine and blunt in
anterior seventh of file length, the other 60—70 sharp. Width of file strongly
increasing in anterior part, reaching maximum 0.35—0.4 mm at end of proximal
fifth of file length, slightly decreasing posteriorly to half maximum width. Spacing
of teeth increasing in anterior seventh of file length, sharp teeth about regularly
spaced. Mirror (pl. 10 fig. 66) elongate, less than twice as long as wide, trapezoidal,
fold much extending over mirror, outline of fold convex, reaching about as far as
postero-apical angle of mirror.

Fore and mid femora with 0—3 spines on anteroventral margin. All knee-lobes
with one spine. Fore and mid tibiae with 1—6 dorsal spines on anterior and 0—2 on
posterior margin, no dorso-apical spines.

Cercus (pl. 9 fig. 56) incurved, tapering apically to a short tooth on inner side of
tip. Subgenital plate (pl. 9 fig. 50) rather three times as long as smallest width, apex
divided by wide triangular incision into slightly narrower triangular lobes with
obtusely pointed tips, the latter representing vestigial styli.

General colour pale and dark brown. Antennae annulated pale and dark brown.
Pronotal dorsum sometimes darker brown, or only laterally so. Fore wing
unicolorous or inconspicuously mottled dark brown, membrane infuscate; hind
margin sometimes narrowly bordered with pale or yellowish brown. Fore and mid
legs of general colour, lower side of fore femur often dark brown. Hind femur of
general colour with lower side between bases of knee-lobes or whole inner lower
margin or whole lower side black. Hind tibia often dark brown from below, except
apically. Spines of tibiae and knees brown, tips black, those of femora usually
completely black. Tarsi from below dull dark brown.

Q (pl. 7 fig. 40), slightly larger than male. Ovipositor very slightly upcurved,
comparatively short, reaching distal end of proximal third of hind tibia and, in
dorsal view, completely covered by flexed wings. Lower margin of ninth tergite
slightly extending over gonangulum (= valvifer), forming shallow furrow. Sub-
genital plate much wider than long, with shallow median apical emargination,
lobes widely rounded. General colour paler brown.

Measurements: body 3 49—53, 9 54—57; fore wing & 57—60, 9 56—64; hind
femur ¢ 34—36, © 34—37; ovipositor 26—30.

Distribution. Known only from the type-locality, East New Guinea, Western
Highlands District: Kandep (map 2).

Discussion. The species is well-defined. This species, together with brevipennis
and stibicki forms the link between Segestes and Segestidea. The thorn-like
fastigium of vertex, the flattened pronotal dorsum and the somewhat truncated

239

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apex of fore wing are found also in Segestidea marmorata and acuminata. No
previous records.

Segestidea I. Bolivar, 1903

Segestidea |. Bolivar, 1903: 166.

Eumossula C. Willemse, 1957: 38 (type-species by monotypy: Eumossula gracilis C. Willemse, 1957).
Syn. nov.

Type-species (by Kirby, 1906: 359): Segestidea princeps 1. Bolivar, 1903
(= Segestidea novaeguineae (Brancsik, 1897)).

Segestidea, as now understood in the key to the genera, is a natural group of
species characterized by the presence of a posterior dorso-apical spine on fore and
mid tibiae (text-fig. 1), in combination with the usual oblique course of Cul of the
fore wing, i.e. vein reaching the hind margin of the fore wing or almost so at short
distance from the wing-base (text-figs. 3, 6, 7).

Up to now, the following species were arranged under Segestidea:
princeps I. Bolivar, 1903
marmorata I. Bolivar, 1903
punctipennis I. Bolivar, 1903
soror Hebard, 1922
hanoverana C. Willemse, 1957
insulana C. Willemse, 1957
The generic distinction among Sexavae as considered in this paper, indicates
that the following taxa also fit in Segestidea:
Sexava femorata C. Willemse, 1940
Sexava leefmansi C. Willemse, 1940
Sexava uniformis C. Willemse, 1940
Segestes acuminatus Kastner, 1934
Sexava rufipalpis C. Willemse, 1966
As will be pointed out, Eumossula C. Willemse, 1957, is synonymous with
Segestidea. As a result, its single species, Eumossula gracilis C. Willemse, 1957,
should be arranged under Segestidea.
Further study of the literature reveals that Moristus novaeguineae Brancsik, 1897,
is synonymous with Segestidea princeps.

For convenience’ sake, a survey is here given of all presently recognized species
and subspecies in Segestidea, noting the new combinations and synonyms:
Segestidea novaeguineae (Brancsik, 1897), comb. nov. (from Moristus) = Segestidea

princeps I. Bolivar, 1903, syn. nov. = Sexava femorata C. Willemse, 1940, syn. nov.

Segestidea punctipennis I. Bolivar, 1903

Segestidea soror Hebard, 1922

Segestidea leefmansi (C. Willemse, 1940), comb. nov. (from Sexava) = Segestidea
hanoverana C. Willemse, 1957, syn. nov.

Segestidea uniformis (C. Willemse, 1940), comb. nov. (from Sexava) = Segestidea

WILLEMSE: Sexavae of Melanesian Subregion 241

insulana C. Willemse, 1957, syn. nov.

Segestidea gracilis gracilis (C. Willemse, 1957), comb. nov. (from Eumossula)
Segestidea gracilis simulatrix ssp. n.

Segestidea rufipalpis (C. Willemse, 1966), comb. nov. (from Sexava)
Segestidea marmorata marmorata I. Bolivar, 1903

Segestidea marmorata occidentalis ssp. n.

Segestidea acuminata (Kästner, 1934), comb. nov. (from Segestes).

Key to the species and subspecies of Segestidea

. Philippine Is. (known only after 9 holotype, body 38 mm, fore wing 29 mm,
ovipositor 21 mm; fore wing with sparse brown dots) . punctipennis I. Bolivar
The Moluccas, New Guinea, Bismarck Archipelago ................. 2
. Fastigium of vertex thorn-like, apex acute, reaching at least apical margin of
SCHADE ET IONI NOIR N TEEN 3
Fastigium of vertex with apex obtusely pointed, shorter, not reaching apical
Maspimorscapeso en N N EN NE emer 5
. Fastigium of vertex reaching pedicel (second antennal segment); fore wing
wider (known only from © holotype from East New Guinea: Milne Bay; may

be synonymous with nominate marmorata I. Bolivar) .. acuminata (Kastner)
Fastigium of vertex reaching first segment of flagellum (third antennal
Segment) fore wine nannower = same es ate ce eet ee ee ne 4

. Male stridulatory file about 3.0 mm long with about 100 teeth (of which 75 are
sharp and apparently functional) (pl. 17 fig. 113) (East New Guinea: Morobe
DIET) ees rere e eee tee eke ee marmorata marmorata I. Bolivar
Male stridulatory file of same length, but with about 200 teeth (of which about
175 are sharp and apparently functional) (pl. 17 fig. 114) (West New Guinea:

Flollandiavareay ee cs ct sae ee ee marmorata occidentalis ssp.n.
. Proximal part of lower and inner side of hind femur solid black or blackish
DOW cape eh O E O AO SOIN Sea ica sh 6
this|partiofhind femur notiblacken waned sara ee 7
. Fore wing with solid blackish brown flecks; knee-lobes with one spine (the
Moluccas: Obi) (known only from the two type specimens) .. soror Hebard

Fore wing unicolorous or slightly mottled dark brown; knee-lobes usually with
two spines (Bismarck Archipelago: New Ireland District) ...............
LIS NER NIE DIL ORE RI MR ER AO: leefmansi (C. Willemse)
. Apical part of hind tibia, over a distance as long as hind tarsus, blackish,
except for upper side; proximal part of outer side of hind femur with a series
of small, black spots, or sometimes with a single large spot, exceptionally
without any spot; ventral spines of hind femur unusually widened basally,
almost triangular (East New Guinea, extending into northeastern part of West
INewiGuinea) Riek LIS AIN PA NE novaeguineae (Brancsik)
Hind tibia unicolorous over whole length; outer side of hind femur always
without solid black spots; ventral spines of hind femur hook-shaped as usual
ALT TER NER 8
. Lower margin of pronotal lateral lobe angulate, deepest point about at distal

242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

third; male stridulatory file as in pl. 16 fig. 108 (East New Guinea: Louisiade

ArchipelagoyMiusimaids); Armani. rufipalpis (C. Willemse)

— Lower margin of pronotal lateral lobe more convex, deepest point about in the

middle; male stridulatory file not as in pl. 16 fig. 108 (Bismarck Archipelago)

EEE A ELLE RITO DIO oo 0 6 cc 9

9. Wings longer, reaching about the middle of hind tibia; ovipositor not

extending beyond apex of flexed wings (pl. 11 fig. 74, pl. 12 figs. 75, 76)

(Admiralty Is. or Manus District) .............. uniformis (C. Willemse)

— Wings shorter, reaching about distal end of proximal fourth of hind tibia;

ovipositor extending far beyond apex of flexed wings (pl. 12 figs. 77—79, pl. 13

figs. 80—82) (New Ireland and New Britain) ...................... 10

10. Anterior part of male stridulatory file narrow, maximum width reached at the
middle of file length (pl. 16 fig. 110) (New Ireland District) ..............

Mine des mes AI AAT RO MAR JE an Rae O gracilis gracilis (C. Willemse)

— Anterior part of male stridulatory file wider, maximum width reached far

before the middle of file length (pl. 16 fig. 111) (East New Britain District)

pis ail dan hal esa nea rete ER ta: Ae etal EE AR gracilis simulatrix ssp.n.

Segestidea novaeguineae (Brancsik, 1897) comb. nov.

(text-figs. 3, 6, 7, pl. 11 figs. 68—71, pl. 15 figs. 91, 99, pl. 16 fig. 107, pl. 17 fig. 115,
map 3)

Moristus novaeguineae Brancsik, 1897: 81.

Sexava novaeguineae; Kirby, 1906: 359.

Sexava coriacea novaeguineae; Karny, 1926: 184.

Segestidea princeps 1. Bolivar, 1903: 167; Kastner, 1934: 53. syn. nov.

Sexava femorata C. Willemse, 1940: 83, figs. 17, 18; 1961: 109, fig. 14; F. Willemse, 1966: 49 (except 19
Manam I.); Lloyd & Gurney, 1975: 47. syn. nov.

Sexava species A: O’Connor, 1959: 122.

Material studied: 9 holotype of Sexava femorata, labelled: Halmaheira, Sexava
femorata n.sp. Det. C. Willemse (CW) (lacking parts of both hind tarsi).

Additional material: Gulf District of Papua, Kerema, ii.1962, sitting on branch of
Coffea canephora (19 CW); Gulf distr., Purari Village, Pawaia no. I., ix—x.1’70, J. I.
Menzies (22 DASF); Port Moresby, viii.1969 (19 CW); Northern District,
Popondetta, Casey’s Plantation, xi.1960, G. S. Dunn, under the bark of bush trees
(19 DASF); Popondetta, season 1964, B. J. Brock (14 19 BMNH); Boana
Mission, Huon Pen., 900 m, 4—5.ix.1956, E. J. Ford Jr., Sexava femorata Will. det.
C. Willemse (1 ¢ BPBM); Bubia via Lae, 12.vii.1960, A. Catley, feeds on coconut
foliage and on Manila hemp foliage (Musa textilis), often parasitized by
Stichotrema dallatorreana Hofeneder (Strepsiptera) (74 99 CW) (partly labelled
Sexava femorata Will., det. C. Willemse); Madang District, Bogia, 1.1960, J. Cox, on
coconut fronds (13 DASF); Manam I., Baliau Village, 26.xi.1960, J. I. Cox,
Sexava femorata Will. det. C. Willemse (24 79 CW); Karkar I., ii.1969, G. R.
Forbes (14 DASF); Karkar I., Kulili Plantation, 20 & 24.ix.1958, J. H. Ardley, ex
coconut palms (44 DASF); Hollandia, viii.1910, Dr. P. N. v. Kampen, Ned. Nw.

WILLEMSE: Sexavae of Melanesian Subregion 243

Guinea Exp. 1911 (44 39 RNH); Hollandia, 24.vii.1938, L. J. Toxopeus, Neth.
Ind. Amer. New Guinea Exp. 1938—39 (1 gu RNH); Maffin Bay, vi.1944, E. S. Ross
(14 19 CAS).

The venation of the fore wing and the presence of fore and mid tibial dorso-
apical spines of the holotype and other material studied of Sexava femorata agree
with Segestidea as here understood. The species is well-defined by a number of
characters, some quite typical. Several of these characters, including the typical
ones, are mentioned in the description of Bolivar’s Segestidea princeps: ‘‘Antennae
— late albido-annulatae — Caput pone oculos fascia pallida obliqua, parum
distincta — Pronotum — lobis deflexis inferne pallidis — Elytra — venis
transversis campi postradialis elevatiusculis, prope venas radiales subcallosus,
flavis — Femora — postica — subtus carnea, extus tertia parte basali maculis fuscis
seriatis ornata, carinis inferioribus — spinis validis armata — Tibiae posticae apice
nigra. Tarsi atri.”. Also the measurements and the locality of princeps (Simbang,
Huon Gulf) are covered by femorata material before me. Unfortunately the
holotype (unique 9) (TMA) has been lost (Steinmann, in litt. 3.xii.1975; V.
Llorente, in litt. 9.1.1976). Moreover, the species was badly known, a second
female only having been recorded by Kastner (1934: 53) who gave no further
comments except for its measurements. Because the typical characters of
Bolivar’s species fit only femorata, both taxa should be considered synonymous.

A similar case offers the comparison of the present material with Brancsik’s
Moristus novaeguineae. That species was known only from the female holotype
from Friedrich Wilhelmshafen (TMA). Again the holotype has been lost
(Steinmann, in litt. 3.xii.1975) and further taxonomically reliable records are not
known. The species was discussed by Karny (1926: 184). Although that author
neiher had the type nor other material before him, he considered it a variety of
Sexava coriacea: ‘‘da sie [novaeguineae] sich von dieser [coriacea] nach der Original
Beschreibung im wesentlichen eigentlich nur durch etwas geringere Dimensionen
und die die Elytren nicht überragende Legeröhre unterscheidet”. However,
Karny’s conclusion is not correct. In Brancsik’s short description it is said: “tibiis
posticis apice tarsisque omnibus subtus fuscus’’. This feature disagrees with any
Sexava species and fits only princeps = femorata. Also the measurements of
novaeguineae agree with the latter rather than with Sexava nubila or coriacea.
Moreover, the type-locality of Brancsik’s species is within the range of princeps =
femorata and not by far within that of Sexava. When comparing the green coloured
specimens before me with Redtenbacher’s monography of the Mecopodinae
(1892: 201), as most probably did Brancsik (1897), the context of the description of
novaeguineae is perfectly clear. Based on so much evidence, I propose to
synonymize Sexava femorata and Segestidea princeps with Moristus novaeguineae.
The last, being the oldest available name, is given priority and the generic
assignment leads to the new combination: Segestidea novaeguineae (Brancsik).

Redescription.
& (pl. 11 fig. 68), large. Fastigium of vertex obtusely pointed, often with shallow
median furrow and slightly fissate apex, reaching to, or slightly extending beyond,

244 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

antennal scrobae, not by far to apical margin of scape. Pron
slightly flattened medially, strongly rounded laterally, anterior margin slightly,
posterior margin less convex to almost straight, “shoulders” indistinct. Pronotal
lateral lobe about as long as high, lower margin angulate, deepest point distally of
the middle of lobe length, this pointsometimes produced posteriorly.

Flexed wings reaching distal end of proximal third to middle of hind tibia. Fore
wing wide, long, margins in proximal half about parallel, beyond middle or in
distal third evenly tapering toward narrowly rounded apex (text-fig. 3); costal area
with numerous parallel obliquely transverse veins, most of these and the transverse
veins of radial and medial areas, slightly incrassate; archedictyon well-developed;
membrane slightly opaque. Venation of bases of left and right fore wing as in text-
figs. 6 and 7. Stridulatory file (pl. 16 fig. 107) fusiform, scarcely arcuate, 4.2—5.0
mm long, number of teeth 175—190 of which anterior 40—50 fine and less sharp,
other 135—150 sharp and apparently functional. Width of file increasing to
maximum 0.5—0.6 mm in anterior third of file length, but slightly decreasing again
posteriorly. Spacing of teeth about regular except for more closely set anterior
ones. Mirror (pl. 17 fig. 115) one-and-a-half times as long as wide or shorter, about
trapezoid; fold strongly inflated in basal half, extending well over mirror, outline
convex basally, running obliquely toward postero-apical angle.

Fore and mid femora usually unarmed, sometimes with 1—3 spines in apical part
of anteroventral margin. Hind femur, except near base, with series of strong
ventral spines on both margins, spines widened basally and almost triangular in
profile. All knee-lobes with two spines, sometimes posterior or seldom also
anterior knee-lobe of mid leg and more often of fore leg, with one spine. Fore tibia
with 0—1, mid tibia with 4—6 dorsal spines on posterior margin, both tibiae with a
posterior dorso-apical spine.

Cercus (pl. 15 fig. 99) incurved, tapering apically to a short, hook-shaped tooth.
Subgenital plate (pl. 15 fig. 91) 5—6 times as long as smallest width, margins about
parallel, apex divided by V-shaped or parabolic incision into pair of lobes, tips of
the latter obliquely truncated and with styli.

General colour bright green or brown. Antennae widely and distinctly annulated
pale yellow and dark brown, except basally. Palpi of general colour, yellowish or
orange. Often whitish fascia from behind eye, extending over genae, lower margin
of pronotal lateral lobe and pleurae. Outer side of hind femur often with pale and
dark transverse stripes, basal third with a series of solid black or dark brown spots,
sometimes fused into one or few larger ones. Lower side of hind femur of general
colour or reddish. Apical part of hind tibia, for a distance about as long as hind
tarsus, black or blackish brown, dorsally usually paler brown or of general colour.
All tarsi from below dull black or blackish brown. Spines of femora black, those of
hind femur usually with basal part reddish brown. Spines of tibiae black, except
for dorsal ones of hind tibia, which are of general colour with tips black.

9 (pl. 11 figs. 69—71) larger than male. Ovipositor straight or almost so, reaching
distal end of proximal third to middle of hind tibia, not or very slightly extending
beyond apex of flexed wings. Subgenital plate wide, triangular, with median apical
emargination and rounded lobes. Coloration as in male.

WILLEMSE: Sexavae of Melanesian Subregion 245

Variation. Measurements vary, but comparative length of wings and ovipositor
are about equal. The contrast between the generally bright green or dark coloured
specimens is conspicuous. The black spots of the basal third of the outer side of
the hind femur are lacking in very few specimens.

Measurements: body ¢ 50—62, 9 56—62; fore wing 3 58— 70, © 75—88; hind
femur 3 39—45, © 46—55; ovipositor 35—42.

Distribution. The range covers the eastern half of the mainland of New Guinea,
extending onto some islands near the coast and into the northern part of western
New Guinea (map 3).

Localities: West New Guinea: Maffin Bay; Takar (O’Connor, 1959); Hollandia.
East New Guinea: East Sepik District: Maprik (O’Connor, 1959); Madang District
(O’Connor, 1959): Friedrich Wilhelmshafen (Brancsik, 1897); Stephansort (Käst-
ner, 1934); Bogia; Manam I. (O’Connor, 1959), Baliau Village (F. Willemse, 1966);
Karkar I. (O’Connor, 1959), Kulili Plantation; Morobe District: Simbang (Bolivar,
1903); Sattelberg; Bubia (F. Willemse, 1966); Boana Mission (C. Willemse, 1961);
Northern District: Popondetta, Casey’s Plantation; Central District: Port Mores-
by; Gulf District: Kerema; Purari Village.

Discussion. The species is well-defined. Easily recognizable are the blackish
apical part of the hind tibia, the black dots on the basal outer side of the hind
femur and the strong spines of the hind femur. The locality label of the holotype of
femorata reads ‘“Halmaheira”, which appears doubtful. The material recorded
under Sexava species A by O’Connor (1959) is not at hand. However, his records
can refer only to novaeguineae and are included in the locality list and on the distri-
bution map.

Segestidea rufipalpis (C. Willemse, 1966) comb. nov.
(pl. 11 figs. 72, 73, pl. 15 figs. 92, 100, pl. 16 fig. 108, pl. 17 fig.116, map 3)
Sexava rufipalpis C. Willemse, 1966: 1, figs. 1, 2; F. Willemse, 1966: 49.

Material studied: type-series, ¢ holo-, 9 allo-, 24 paratypes, labelled: Liak
Village, Misima Island, Milne Bay District, of Papua, Feb. 1962; feeding on the
foliage of Cocos nucifera, coll. D. I. Murrie, Sexava rufipalpis sp.n. Det. C.
Willemse 1962, appropriate type-labels (CW) (holotype lacks right antenna, right
fore leg, both mid tibiae and tips of both hind tarsi).

For a general description, compare original one. Differs from the type-species
as follows.

More robust but not larger (pl. 11 figs. 72, 73). Apex of flexed wings not reach-
ing beyond proximal third of hind tibia and almost beyond tip of ovipositor. Fore
wing with apex narrowly rounded, almost pointed. Male stridulatory file (pl. 16 fig.
108) distinctly fusiform, almost straight, about 4 mm long, number of teeth
95—105 of which anterior 15—20 very fine, blunt, closely set and weakly
sclerotized, arranged over anterior ninth of file length, the other 80—90 sharp and
strongly sclerotized. Width of file strongly increasing in anterior ninth, from there

246 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

but slightly and reaching maximum of 0.5 mm at distal end of anterior fourth of
file length, decreasing posteriorly to about one-fourth of maximum width. Spacing
of nine anterior teeth very close, of remaining part of file about regular and much
more widely spaced than in type-species. Mirror (pl. 17 fig. 116) less trapezoid,
more elongate-elliptical, fold extending less over mirror and less inflated. Spines of
hind femur not widened basally but hook-like as usual. Male cercus and subgenital
plate as in pl. 15 figs. 92, 100.

General colour green. Antennae scarcely annulated. Palpi orange. Whitish
fascia of head and thorax scarcely indicated. Pronotal dorsum, on each side, with a
minute black point in the impressed posterior transverse sulcus. Legs, including
apical part of hind tibia and outer side of hind femur and tarsi, of general colour.
Dorsal spines of hind tibia orange brown, tips black, other spines of legs
completely black or almost so.

Measurements: body 3 47—52, 9 60; fore wing & 57—64, © 70; hind femur ¢
39— 40, © 48; ovipositor 39.

Distribution. Known only from the type-locality, Papua, Milne Bay District:
Louisiade Archipelago, Misima I. (map 3).

Discussion. The species is well-defined. The male stridulatory file is quite
characteristic. The shape of the pronotal lateral lobe and the male abdominal
terminalia are much as in novaeguineae.

Segestidea uniformis (C. Willemse, 1940) comb. nov.

(pl. 11 fig. 74, pl. 12 figs. 75, 76, pl. 15 figs. 93, 101, pl. 16 fig. 109, pl. 18 fig. 117,
map 3)

Sexava uniformis C. Willemse, 1940: 81, figs. 19, 20 (only Lou I.).
Segestidea insulana C. Willemse, 1957: 41, pl. 4 right; 1961: 111 (only Los Negrlitlos); F. Willemse, 1966:
48 (only paratypes of insulana and Wululu I.). syn. nov.

Material studied: 9 holotype of Sexava uniformis, labelled: Lou Il. ’32, coll.
Bohler, Sexava n. sp. det. Karny, Sexava uniformis nov. sp. Det. C. Willemse, Type
(NMB) (discoloured, both fore legs, right middle leg and left fore wing lacking).

Type-series of Segestidea insulana: & holotype, labelled: New Guinea Territory
of Papua, Pak Island, 1954, T. H. Ardley, Segestidea insulana nov. spec. det. C.
Willemse, type (BMNH); paratypes: similar locality label (14 19 BMNH; 19
CW); Lou I., 1954, T. H. Ardley (14 19 BMNH); Lorengau Manus I., 1954, T. H.
Ardley (1 ¢ CW; 19 BMNH) (with appropriate identification and type-labels).

Additional material: Manus District, 24—28.ii.1974, J. Pippett (14 DASF);
Manus District, Sala Plantation, 4.iii.1954, J. Ardley (19 DASF); Manus I,
5.v.1932, J. L. F. (13 29 BMNH); Manus, 1932, N. E. H. Caldwell (14 BMNH);
Manus I., Lei Village near Lorengau, 29.vi.1956, J. Szent-Ivany, feeding on fronds
of coconut palms (14 19 DASF); Manus I., Tulo Plantation N. coast, 19.1.1962,
on Cocos nucifera, J. Szent-Ivany & P. Hermann (3¢ DASF); Manus I., Bundalis
R. C. Mission, plantation N.coast, 19.1.1962, on mature coconuts, brought down
by smoke, J. Szent-Ivany & P. Hermann (24 DASF); Los Negros Is., Momote

WILLEMSE: Sexavae of Melanesian Subregion 247

Airstrip, 26.11.1960, defol. young coconut palms, J. Szent-Ivany (14 DASF); Los
Negros, xi.1945, Wagner & Grether, Segestidea insulana Will. det. C. Willemse (14
BPBM); Pak I., 7.1.1971, P. R. Jones, on coconuts (34 49 DASF); Hermit Is.,
Maron I., 10.vii.1961, ex coconut palms, J. H. Ardley (Sg 59 CW); Wululu Isl.,
Agita Plantation, ix.1960, Rede Lean, severe damage to coconut foliage,
Segestidea insulana Will. det. C. Willemse (6¢ 19 CW).

Sexava uniformis was described after two females, the holotype from Lou I., the
paratype from Rook I. The latter represents another species and is discussed
under gracilis simulatrix. Comparison of the holotype of uniformis, the type-series
of insulana and other material at hand reveals clearly that they belong to one
species, to be assigned to Segestidea.

For a general description the reader is referred to the original descriptions of
Sexava uniformis and Segestidea insulana. The species differs from the type-species
as follows.

Smaller, more slender (pl. 11 fig. 74, pl. 12 figs. 75, 76). Pronotal lateral lobe
narrower, lower margin obtusely angulate, deepest point about in the middle.
Flexed wings reaching middle of hind tibia or almost so, in dorsal view covering
Ovipositor completely. Ovipositor straight, tip reaching distal end of proximal third
of hind tibia or slightly longer. Fore wing much narrower, comparatively less
tapering toward comparatively more widely rounded apex; archedictyon less well-
developed; transverse veins in costal area less numerous and less regular, not
incrassate; membrane more or less transparent. Male stridulatory file (pl. 16 fig.
109) fusiform, slightly arcuate, 3.4—4.0 mm long, number of teeth 130—140, of
which anterior 35—50 fine, less sharp, and others (at least 85—95), sharp and
apparently functional. Width of file increasing and reaching maximum 0.48—0.50
mm at distal end of anterior third of file length, slightly decreasing again
posteriorly to about half maximum width. Spacing of teeth increasing in anterior
third, functional teeth about regularly set. Mirror (pl. 18 fig. 117) about twice as
long as wide, elongate-elliptical, fold distinctly less extending over mirror and but
weakly inflated, outline straight and about parallel to hind margin of wing.
Number of spines on legs about as in type-species, ventral spines of hind femur not
widened, but hook-like as usual. Male cercus shorter (pl. 15 fig. 101) and very
slightly incurved apically. Male subgenital plate (pl. 15 fig. 93) narrow, lateral
margins often slightly concave, margins of V-shaped apical incision often slightly
crenulated, styli comparatively longer and slightly incurved.

General colour green or brown. Antennae not or scarcely annulated. Head,
thorax, wings and all legs of general colour. Fore wing sometimes with hind
margin narrowly dark brown or yellowish. Spines of legs of general colour, tips
black. Tarsi from below dull dark brown.

Measurements: 4 43—52, 9 42—57; fore wing 4 57—65, 9 62—74; hind femur
Sd 32—41, 9 34—44; ovipositor 26—33.

Distribution. As far as known, the range covers the north-western Bismarck
Archipelago: Admiralty Is. (Manus District) (map 3).

Localities: Manus I.: Lorengau (C. Willemse, 1957); Lei Village; Tulo Planta-
tion; Bundalis R. C. Mission; Sala Plantation; Lou I. (C. Willemse, 1940; 1957);

248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

Pak I. (C. Willemse, 1957); Los Negros I. (C. Willemse, 1961), Momote Airstrip;
Hermit Is., Maron I.; Wululu Is., Agita Plantation (C. Willemse, 1966).

Discussion. The species is well-defined, but the characters are not as easily
recognizable as in some other species of the genus. Confusion is possible with
nominate Segestidea gracilis and its subspecies simulatrix, and with Segestes
decoratus. Distinction from the latter is discussed under that species. Both
subspecies of gracilis have shorter wings and longer ovipositor, the latter extending
far beyond the apex of the flexed wings (pl. 12 figs. 77—79, pl. 13 figs. 80—82).
Moreover the lower margin of the pronotal lateral lobe is more convex, the
male cercus more incurved (pl. 15 figs. 102, 103), the male subgenital plate wider
(pl. 15 figs. 94, 95), and the styli shorter. The male stridulatory file in nominate
gracilis has a quite different shape (pl. 16 fig. 110), while the file in simulatrix is less
arcuate and differs in slightly wider spacing of the teeth (pl. 16 fig. 111).

Segestidea gracilis (C. Willemse, 1957) comb. nov.

Among the material before me, two taxa can be recognized, based on consistent
differences in the stridulatory file, as indicated in the key above. This may be
evidence of genetic discontinuity in which case the taxa should be considered
species. However, by lack of more supporting evidence, I prefer to consider them
subspecies. One occurs in New Ireland and agrees with gracilis, the other
subspecies is known from New Britain and named simulatrix ssp.n.

Segestidea gracilis gracilis (C. Willemse, 1957) comb. nov.

(pl. 12 figs. 77—79, pl. 15 figs. 94, 102, pl. 16 fig. 110, pl. 18 fig. 118, map 3)

Eumossula gracilis C. Willemse, 1957: 37, figs. 1—3, pl. 2; 1958: 122 (only New Ireland); F. Willemse,
1966: 48 (do.).
Segestidea insulana; C. Willemse, 1961: 110 (only New Ireland); F. Willemse, 1966: 48 (do.).

Material studied: type-series of Eumossula gracilis, 3 holo-, 14 19 paratypes,
labelled: New Guinea, Terr. of Papua, New Ireland, 1954, J. H. Ardley, Eumossula
gracilis n.g. n.sp. Det. C. Willemse, appropriate type-labels (holo-, 14 paratype
BMNH; 19 CW) (holotype lacks right antenna and some claws).

Additional material (all New Ireland District): Lihir I., Londolovit Plantation,
on coconut fronds, 20.vii.1955, J. Szent-Ivany (3g CW); Lihir Group, Masahet I.,
on coconut palms, 21.vii.1955, J. Szent-Ivany (24 29 CW); Lihir Group, Mahur
I., on coconut palms, 22.vii.1955, J. Szent-Ivany (14 29 CW); SW New Ireland,
Gilingil Plantation, 2 m, 17.vii.1956, J. L. Gressitt (1 ¢ CW; 13 BPBM) (part of all
this material with label: Segestidea insulana Will. Det. C. Willemse); New Ireland,
Tigak D.A.S.F., 1. viii.1971, on coconut, J. Sumbak (19 DASF); Tabar I., Teripax
Plantation, 23.vii.1955, on Cocos nucifera, J. Szent-Ivany (1 ¢ DASF); Lihir Group,
Lataul Village, 15.xi.1968, D. F. O’Sullivan (14 DASF); New Ireland District,
Namatanai, Matakan Plantation, 15.iii.1965, on Theobroma cacao, J. M. Adams

WILLEMSE: Sexavae of Melanesian Subregion 249

(1g 19 DASF); New Ireland District, Maramakas Plantation, 7.11.1954, J. H.
Ardley (13 DASF).

A study of the type-series and other available material reveals clearly that the
monotypic Eumossula perfectly fits Segestidea and that its species, gracilis, is well-
defined.

For a general description the reader is referred to the original one. The species
differs from the type-species as follows.

Smaller (pl. 12 figs. 77—79). Lower margin of pronotal latera lobe slightly
convex, not at all angulate. Wings shorter, in flexed position extending just beyond
hind knee, not beyond proximal fourth of hind tibia, not by far reaching tip of
ovipositor. Fore wing much narrower, slightly tapering toward narrowly rounded
apex; archedictyon well-developed; transverse veins less numerous, less regular
and not incrassate. Male stridulatory file (pl. 16 fig. 110) narrowly fusiform, slightly
arcuate, 3.8—4.2 mm long, number of teeth 156—175 of which at least 50— 70
cover the atenuated, narrow anterior third of file length, while others (about 100)
are arranged over remainder of file. Width of file increasing but slightly all over
anterior half, just before or in middle of file length increasing more abruptly,
reaching maximum of 0.4—0.5 mm and decreasing again slightly in posterior
fourth of file. Spacing of teeth slightly and gradually increasing in anterior half,
from there slightly decreasing posteriorly. Mirror (pl. 18 fig. 118) wide elliptical,
fold not strongly inflated, outline of the latter almost straight of slightly sinuate
and but moderately extending over mirror. Number of spines of legs as in type-
species. Ventral spines of hind femur hook-like, as usual. Male cercus (pl. 15 fig.
102) shorter, more robust. Male subgenital plate (pl. 15 fig. 94) comparatively
shorter, about four times as long as smallest width. Ovipositor long, reaching
middle of hind tibia or almost so, upper margin straight.

General colour pale green or brown. Antennae not or scarcely annulated. Head,
thorax, fore wings and legs of general colour, spines with tips black.

Measurements: body ¢ 47—54, 9 48—59; fore wing ¢ 48—54, 9 54—63; hind
femur ¢ 37—42, © 40—44; ovipositor 36—40.

Disribution. The range of the nominate subspecies covers the New Ireland
District of the Bismarck Archipelago (map 3).

Localities: New Ireland (C. Willemse, 1957); Giingil Plantation (C. Willemse,
1961; F. Willemse, 1966); Maramakas Plantation; Namatanai, Matakan Planta-
tion; Tigak; Lihir Is., Lataul Village; Londolovit Plantation (C. Willemse, 1958; F.
Willemse, 1966); Masahet I. (C. Willemse, 1958; F. Willemse, 1966); Mahur I. (C.
Willemse, 1958; F. Willemse, 1966); Tabar I., Teripax Plantation.

Discussion. The nominate subspecies is characterized by short wings, long
ovipositor and the shape of the male stridulatory file. As to the latter, it is noted
that this character is rather uniform throughout the studied males of nine different
localities.

Distinction with Segestidea uniformis and Segestes decoratus is discussed under
these species.

250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

Segestidea gracills simulatrix subsp. nov.

(pl. 13 figs. 80—82, pl. 15 figs. 95, 103, pl. 16 fig. 111, pl. 18 fig. 119, map 3)

Sexava uniformis C. Willemse, 1940: 82, figs. 19—20 (Rook I. only).
Segestidea insulana; C. Willemse, 1961: 110 (New Britain only); F. Willemse, 1966: 48 (do.).
Eumossula gracilis; C. Willemse, 1958: 122 (New Britain only); F. Willemse, 1966: 48 (do.).

Material studied: ¢ holotype, labelled: New Britain, Gazelle Peninsula, Lowl.
Agr. Exp. Station Keravat, 26.v.1954, J. Szent-Ivany, on the fronds of African oil-
palms Elea guineensis, Eumossula gracilis Will. Det. C. Willemse 1958 (CW) (both
antennae, left fore tibia and parts of tarsi lacking).

Paratypes: similar to holotype (1 9 CW), similar locality and identification label,
viii.1958, G. S. Dunn, on Cocos nucifera (13 29 CW), similar locality, 60 m,
11.1x.1955, J. L. Gressitt, Segestidea insulana Will. Det. C. Willemse (14 29
BPBM; 19 CW); New Britain, Gazelle Peninsula, Bainings St. Paul’s, 350 m, 9. ix.
1955, J. L. Gressitt, Segestidea insulana Will. Det. C. Willemse (134 49 BPBM;
1g CW); New Britain, Gazelle Peninsula, Malaguna, 4.iii.1971, feeding on foliage
of Cocos nucifera, A. Gameta & J. Guguna (3¢ 29 DASF); New Britain, Gaulim
Peninsula, 23.v.1955, Segestidea insulana Will. det. C. Willemse (19 BPBM); New
Britain, Sio, N. coast, 600 m, 24.vii.1956, Segestidea insulana Will. det. C. Willemse
(13 BPBM); New Britain, Ti, Nakagai, 28.vii.1956, E. J. Ford, Segestidea insulana
Will. det. C. Willemse (14 BPBM); E. N. B. District, 30.xii.1974, on coconut leaf,
D. Tago (19 DASF); New Britain, Kokope East, Gunanua Plantation, 27.iv. 1968,
on oil palm, R. Abijah (19 DASF); East New Britain, Sumsum Plantation,
31.111.1971, Cocos nucifera, D. F. O’Sullivan (19 DASF); Rook I., Umboi, 1930, H.
Hediger, Sexava uniformis nov. spec. Det. C. Willemse, cotype (19 NMB).

Description.

Differing from nominate subspecies as follows (pl. 13 figs. 80—82). Male
stridulatory file (pl. 16 fig. 111) slightly fusiform, scarcely arcuate, 3.8—4.2 mm
long, number of teeth 117—139 of which anterior 20—40 fine and less sharp
covering anterior fifth of file length, other teeth, about 100, arranged over
remaining of file. Width of file stronger, increasing in anterior fifth, reaching
maximum of 0.4—0.5 mm at distal end of proximal third of file length, from there
slightly decreasing posteriorly. Spacing of teeth increasing in anterior third, about
regular in remaining of file. Mirror (pl. 18 fig. 119) slightly more elongate. Male
cercus (pl. 15 fig. 103) slightly more incurved.

General colour green or brown, sometimes slightly mottled darker brown,
especially in fore wing.

Measurements: body ¢ 46—52, 9 53—59; forewing g 45—50, 9 53—58; hind
femur ¢ 32—36, 2 38—40; ovipositor 36—38.

Distribution. As far as known confined to New Britain (and Rook I. ?) (map 3).

Localities: East New Britain: Sumsum Plantation; Malaguna; Kokopo, Gunanua
Plantation; Bainings St. Paul’s (C. Willemse, 1961; F. Willemse, 1966); Keravat

WILLEMSE: Sexavae of Melanesian Subregion 251

Lowlands Agric. Exper. Station (C. Willemse, 1958, 1961; F. Willemse, 1966); New
Britain: Ti, Nakagai; Sio, N. coast; Gaulim Peninsula; Rook I.

Discussion. This subspecies is characterized by its resemblance with the
nominate subspecies in combination with its clearly distinct male stridulatory file.
As in the nominate subspecies, it is noted that the file of the studied males from
five different localities is quite uniform. Whether the slight differences of the male
cercus, subgenital plate and mirror (pl. 15 figs. 95, 103, pl. 18 fig. 119) are reliable
characters, is not certain by lack of sufficient material. The paratypic Sexava
uniformis female from Rook I. is discoloured and badly damaged. Without the
male, identification is uncertain, although it can be said that the specimen
disagrees with uniformis and Segestes decoratus. Judging from the length of the
wings and the ovipositor and the shape of the pronotal lateral lobe, it fits gracilis.
As to the range of the subspecies of the latter, simulatrix may be expected to occur
in Rook I. rather than the nominate form.

Distinction between gracilis simulatrix, Segestidea uniformis and Segestes de-
coratus is discussed under the last two species.

Segestidea leefmansi (C. Willemse, 1940) comb. nov.

(pl. 13 figs. 83, 84, pl. 14 fig. 85, pl. 15 figs. 96, 104, pl. 16 fig. 112, pl. 18 fig. 120,
map 3)

Sexava leefmansi C. Willemse, 1940: 83, fig. 21—22.
Segestidea hanoverana C. Willemse, 1957: 39, pl. 3, 4 left; F. Willemse, 1966: 48. syn. nov.

Material studied: type-series of Sexava leefmansi: 3 holotype, labelled: La-
wongai N.H. xi.31 coll. Buhler, Sexava sp. n. det. H. Karny, Sexava leefmansi nov.
sp. adet. C. Willemse, type (discoloured, left fore leg and right fore tibia lacking);
paratypes: similar labels, cotype (14 19); Namaoroso N.H. xi.31 coll. Buhler,
similar identification labels, cotype (14 19) (all NMB).

Type-series of Segestidea hanoverana: 3 holo-, 24 39 paratypes, labelled: New
Guinea, Terr. of Papua, New Hanover, 1954, J. H. Ardley, Segestidea hanoverana
nov. sp. Det. C. Willemse, appropriate type-labels (holotype lacks left mid leg,
BMNH;1g 29 BMNH; 1 4 19 CW).

Additional material (all New Ireland District): Feni Island Group, Anir
Plantation, on Cocos nucifera, x.1959, G. S. Dunn (19 19 CW); Tabar Group,
Tatau Island, Teripax Plantation, defoliating coconuts, severe outbreak, 23.vii.
1955, J. Szent-Ivany (24 1 9 CW; 1g DASF); New Ireland Eastcoast, Metakabul
Plantation, on coconuts, 21.viii.1955, J. Szent-Ivany (14 19 CW) (all labelled
Segestidea hanoverana Will. Det. C. Willemse); New Hanover, Umbukul, 15.111.
1971, resting on fronds of Cocos nucifera, B. Dionsil (29 DASF); Lihir Group,
Masahet Island, 21.ix.1955, J. Szent-Ivany, on old coconut palm (1 ¢ DASF); Lihir
Group, Mahur Island, 21.vii.1955, on young palm, J. Szent-Ivany (13 DASF);
New Hanover, Baikep Village garden, 21.viii.1955, J. Szent-Ivany, on young

252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

coconut palms (19 DASF); New Hanover, Tioputuk & Metevol, 11—15.111.1971,
B. Dionsil, resting on fronds of Cocos nucifera (19 DASF); New Hanover,
5.v.1932, coconuts (13 19 BMNH).

Comparison of the types of Sexava leefmansi, Segestidea hanoverana and other
material at hand reveals clearly that they are conspecific and should be arranged
under Segestidea.

A general description can be found in the original ones of leefmansi and
hanoverana. The species differs from the type-species as follows.

Slightly smaller (pl. 13 figs. 83, 84, pl. 14 fig. 85). Lower margin of pronotal
lateral lobe from regularly to obtuse-angularly rounded, deepest point about in
middle. Flexed wings reaching from distal end of proximal third to middle of hind
tibia, in female to tip or just beyond tip of ovipositor. Fore wings long and wide,
margins about parallel, in apical third hind margin more and fore margin less
tapering towards widely rounded apex; archedictyon well-developed; transverse
veins less numerous, in costal area irregular. Male stridulatory file (pl. 16 fig. 112)
narrowly fusiform, slightly arcuate, 3.0—3.2 mm long, number of teeth 132—145
of which about 50—70 cover anterior third of file length. Width of file but very
slightly and gradually increasing in anterior third of file length, reaching maximum
of 0.3 mm near middle of file, from there slightly decreasing posteriorly. Spacing
of teeth slightly and gradually increasing in anterior third, about regular over
remaining of file length. Mirror (pl. 18 fig. 120) elongate-elliptical, about 3 mm
long and 2 mm wide, fold moderately developed and inflated, its outline convex to
slightly sinuate. Spines of legs as in type-species, ventral ones of hind femur hook-
like as usual. Male cercus (pl. 15 fig. 104) much less incurved, near apex abruptly
narrowing to short, strong, incurved tooth. Male subgenital plate (pl. 15 fig. 96)
narrow, 6—7 times as long as smallest width, slightly widening apically, lateral
margins slightly concave, margins of apical incision often crenulated. Ovipositor
with upper margin straight or almost so, reaching from distal end of proximal third
to middle of hind tibia.

General colour from pale to dark brown, seldom green. Antennae slightly
annulated pale and dark brown. Head and thorax of general colour. Fore wing
usually mottled with darker brown areas and dots, membrane infuscated. Legs of
general colour, basal part of lower and inner side of hind femur solid black. Spines
brown, tips black, tarsi from below dull dark brown.

Measurements: body ¢ 46—53, 9 51—59; fore wing 4 59—65, 9 72—75; hind
femur G 40—42, © 41—45; ovipositor 30—34.

Distribution. As far as known, confined to the New Ireland District of the
Bismarck Archipelago (map 3).

Localities: New Hanover (C. Willemse, 1957); Lawongai (C. Willemse, 1940);
Namaoroso (C. Willemse, 1940); Tioputuk and Metevol; Umbukul; Baikep; New
Ireland, Metakabul Plantation (F. Willemse, 1966); Feni Is., Anir Plantation (F.
Willemse, 1966); Tabar Is., Tatau I., Teripax Plantation (F. Willemse, 1966); Lihir
Group, Masahet I.; Mahur I.

WILLEMSE: Sexavae of Melanesian Subregion 253

Discussion. The species is well-defined by a number of characters. The
coloration of the hind femur is a conspicuous feature. The male stridulatory file
resembles that of gracilis simulatrix, but is shorter and narrower.

In the original description of leefmansi, the type-localities were recorded from
the New Hebrides. Apparently this was a mis-interpretation for the abbreviation
N.H. on the locality-label. Direct information given by Prof. Bühler, who
collected the specimens (via Dr. Baroni-Urbani, in litt. 24.x1.1975) confirms that
the specimens originate from New Hanover. As far as known, Sexavae do not
occur inthe New Hebrides.

The species lives together with Segestidea gracilis gracilis.

Segestidea acuminata (Kästner, 1934) comb. nov.

(pl. 14 figs. 86, 87, map 3)
Segestes acuminatus Kästner, 1934: 50, figs. 18—19.

Material studied: 9 holotype, labelled: Coll. Br. v.W. Milne Bay (Neu Guinea)
Staud., Segestes sp. determ. Karny, 21.673, Segestes acuminatus Kast. 9 A. Kästner
det., Typus (NMW) (both antennae, right fore tibia, tarsi of four anterior legs,
both cerci, tip of right middle tibia lacking and tips of both fore wings slightly
damaged).

The holotype (pl. 14 figs. 86, 87) agrees fairly well with the original description
except for two details. The intact left fore and mid tibiae bear a posterior dorso-
apical spike, both apparently overlooked by Karny who studied the specimen, and
by Kastner when describing his species. This character disagrees with Segestes and
assigns the species to Segestidea. Further it can be seen that the apex of the fore
wing, although slightly damaged, is truncated rather than narrowly rounded.

Other marked characters are the thorn-like fastigium of the vertex, which
extends just beyond the scape. The pronotal dorsum is somewhat flattened in the
middle, slightly rounded laterally, fore and hind margins each bearing two lateral
and one median obtuse tubercles. Lower margin of pronotal lateral lobe obtuse-
angularly rounded, deepest point just before the middle. The flexed wings reach
the tip of the ovipositor, which reaches as far as the middle of the hind tibia. The
venation of the fore wing agrees with that of the genus. Ovipositor is compara-
tively short and slightly upcurved. The subgenital plate is wider than long, slightly
incised apically with widely rounded lobes. Fore and mid femora with 5—6 spines
on the anteroventral margin. All knee-lobes with one spine. Fore and mid tibiae
with 3 dorsal spines on the anterior margin, posterior margin of fore tibia
unarmed, that of the mid tibia with 3 spines, dorso-apical spines not included.
General colour is rusty brown. The apex of the fastigium of the vertex and a lateral
stripe over the pronotal dorsum blackish. Fore wing with scattered dark brown
points, caused by dark brown membrane between the archedictyon.

Measurements: body ¢ 38; fore wing 47; hind femur 28; ovipositor 20.

Distribution. Known only from the holotype, Papua: Milne Bay (map 3).

Discussion. The species is badly defined. It comes near marmorata (see below).

254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

The distinction indicated in the key appears not reliable. Topotypic material,
including the male, is needed to establish the differences between both taxa.

Segestidea marmorata I. Bolivar, 1903

On account of similar considerations as given for gracilis, two subspecies in
marmorata can be recognized: nominate marmorata and marmorata occidentalis
ssp.n.

Segestidea marmorata marmorata I. Bolivar, 1903
(pl. 15 figs. 97, 105, pl. 17 fig. 113, pl. 18 fig. 121, map 3)

Segestidea marmorata I. Bolivar, 1903: 167.
Segestes acuminatus; C. Willemse, 1961: 108, figs. 18, 19.

Material studied: E. New Guinea, Bubia, Markham Valley, 50 m, 19.ix.1955, J.
L. Gressitt, Segestes acuminatus Kastner Det. C. Willemse 1960 (1 ¢ BPBM).

The species was described after a single female from: Sattelberg, Golfe Huon,
Bird, 1898. Until now, no further material became available. Unfortunately the
holotype has been lost (TMA) (Steinmann, in litt. 3.xii.1975) and other material
could not be traced (IEM; V. Llorente, in litt. 9.i.1976). The male before me agrees
fairly well with Bolivar’s description and its locality is not far from the type-
locality. Preliminarily, the specimen is assigned to nominate marmorata. It differs
from the type-species as follows.

Redescription.

d smaller, more slender (C. Willemse, 1961: fig. 19). Fastigium of vertex (C.
Willemse, 1961: fig. 18) thorn-like, apex acute, slightly upcurved, extending far
beyond the antennal scrobae, reaching apical margin of scape. Pronotal dorsum
somewhat flattened medially, slightly rounded laterally, anterior margin almost,
posterior margin quite straight, both margins with weak median tubercle. Pronotal
lateral lobe about as long as high, lower margin obtusely angulate, deepest point
about in the middle.

Flexed wings reaching just beyond middle of hind tibia. Fore wing long and
narrow, margins about parallel, very slightly narrowing towards wide, obliquely
truncated and slightly emarginate apex; archedictyon well-developed; transverse
veins not incrassate, ill-defined and but few in number. Stridulatory file of the
male at hand (pl. 17 fig. 113) partly torn off from the membrane, fusiform, slightly
arcuate, 2.9 mm long, number of teeth 105 of which anterior 30 fine and about
blunt, covering anterior seventh of file length, remaining 75 teeth sharp. Width of
file strongly increasing in anterior part, reaching maximum 0.4 mm in middle of
file, decreasing posteriorly. Spacing of teeth very narrow in anterior part, strongly
increasing towards middle of file, from there about regular and scarcely
decreasing posteriorly. Mirror (pl. 18 fig. 121) roughly trapezoid, about 3 mm long
and 2 mm wide, fold extending well over mirror with strongly inflated basal half,

WILLEMSE: Sexavae of Melanesian Subregion 255

its outline convex basally, running obliquely towards postero-apical angle of
mirror.

Fore and mid femora with 3 spines on apical half of anteroventral margin. All
knee-lobes with one spine. Fore tibia with 3 and 6, mid tibia with 2 and 3 dorsal
spines on posterior and anterior margins, respectively, dorso-apical spines not
included. Spines of hind femur hook-like. Cercus (pl. 15 fig. 105) slender, incurved
and slightly tapering apically. Subgenital plate (pl. 15 fig. 97) small, narrow, about
four times as long as smallest width, apex divided by V-shaped incision into pair of
narrow lobes, styli present.

General coloration mottled pale and dark brown. Fastigium of vertex from
above, pronotal dorsum and cubito-anal areas of fore wing, dark brown. Antennae
slightly annulated. Fore wing distinctly mottled pale and dark brown, combined
with smaller and larger areas of transparent membrane; hind margin narrowly
bordered yellowish white in proximal half. Inner side of hind femur castaneous
brown. Spines of legs black, dorsal spines of hind tibia brown, tips black.

Q, after Bolivar: Lamina subgenitalis postice rotundata, medio obtuse angulato-
emarginata. Ovipositor subrectus, pallidus, apice magis infuscatus.

Measurements (9 after Bolivar): body & 46, © 45; fore wing ¢ 59, © 59; hind
femur ¢ 31, 9 33; ovipositor 24.

Distribution. Known only from East New Guinea: Morobe District (map 3).

Localities: Sattelberg (Bolivar, 1903); Bubia.

Discussion. The nominate subspecies is well-defined, although variation is
insufficiently known. It links Segestidea with Segestes, which is apparent when
comparing marmorata with Segestes stibicki, brevipennis and cornelii. As pointed out
under acuminata, that species might be synonymous with nominate marmorata.

Segestidea marmorata occidentalis subsp. nov.

(pl. 14 figs. 88 —90, pl. 15 figs. 98, 106, pl. 17 fig. 114, pl. 18 fig. 122, map 3)

Material studied. ¢ holotype, labelled: Neth. New Guinea, Dojo, iv.1958, G.
den Hoedt (CW); paratypes: Hollandia, Ned. Nieuw Guinea Exp. 1911 Dr. P.N. v.
Kampen (23 29 RNH) (holotype lacks both antennae and part of some tarsi,
paratypes discoloured and more damaged).

Description.

d (pl. 14 figs. 88, 89). Differs from nominate subspecies in larger number of
teeth of male stridulatory file (pl. 17 fig. 114). The latter of similar length, 2.9—3.1
mm, number of teeth about twice as large, 198—203, maximum width of file
slightly smaller, 0.3 mm, spacing of teeth twice as narrow. Mirror (pl. 18 fig. 122)
of right male fore wing, abdominal terminalia (pl. 15 figs. 98, 106) and coloration
about as in nominate subspecies.

Q (pl. 14 fig. 90). Flexed wings reach middle of hind tibia, extending beyond tip
of ovipositor. Ovipositor short, slightly upcurved, reaching about distal end of
proximal third of hind tibia. General coloration more uniform.

256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

Measurements: body ¢ 39—47, 9 36—40; fore wing & 56—57, g 51—52; hind
femur & 31—33, Q 30—31; ovipositor 20—21.

Distribution. Known only from the type-series, West New Guinea: Hollandia
and nearby Dojo (map 3).

Discussion. More material is needed to establish precisely the distinction
between occidentalis and nominate marmorata. Although the number of teeth of
the male stridulatory file in Sexava coriacea varies in a similar degree, that number
is defined by the length of the file, while the spacing of the teeth is about similar.
This is not the case in the marmorata material, which justifies the subspecific
distinction between the nominate form and occidentalis.

No previous records.

Segestidea soror Hebard, 1922
Segestidea soror Hebard, 1922: 178, pl. 16 figs. 3, 4.

Known only from the typical pair (ANSP).

Distribution. The Moluccas: Obi (Hebard, 1922).

Discussion. I have before me a male with the tip of abdomen badly damaged.
It is labelled: Dodinga, Halmaheira, Bernstein (CW). As far as can be judged, the
specimen agrees with Hebard’s species. Especially the shape of the fore wing with
obliquely truncated apex fits soror perfectly. The stridulatory file is 3.9 mm long,
slightly arcuated and fusiform, number of teeth 79 of which the anterior 28 fine
and less sharp than the remaining 51 ones. Width of the file reaching maximum
of 0.47 mm in anterior fourth of file length, decreasing posteriorly to about a third
of the maximum width. Spacing of the teeth very fine and increasing in anterior
third, decreasing posteriorly to about half widest spacing. Mirror twice as long as
wide, elliptical; fold extending well over the mirror, outline almost straight and
parallel to the hind margin of the wing.

Segestidea punctipennis I. Bolivar, 1903

Segestidea punctipennis 1. Bolivar, 1903: 168.

Known only from the 9 holotype (IEM).
Distribution. The Philippine Is.: Irocin, Albay (Bolivar, 1903).

REFERENCES

Beier, M., 1955. Emboidea und Orthopteroidea. — In: H. G. Bronn’s Klassen und Ordnungen des Tier-
reichs 5 (3) 6: 1—304, figs.

———, 1962. Orthoptera Tettigoniidae (Pseudophyllinae I). — Das Tierreich 73: 1—468, figs.

———, 1966. Tettigoniidae: Subfam. Meconematinae, Mecopodinae, Phyllophorinae. — In: M. Beier,
Orthopterorum Catalogus 9: 248—342.

Brancsik, C., 1897. Series orthopterorum novorum. — Jh. naturw. Ver. Trencsiner Kom. 19—20
(1896— 1897): 52—85, figs.

WILLEMSE: Sexavae of Melanesian Subregion 25

Bolivar, I., 1903. Contributions a l’étude des Mecopodinae. — Annls Mus. nat. hung. 1: 161—178.

Brunner von Wattenwyl, C., 1878. Monographie der Phaneropteriden. — K.K. zool. -bot. Ges. Wien:
1—401, figs.

———, 1898. Orthopteren des Malayischen Archipels gesammelt von Prof. Dr. W. Kükenthal in den
Jahren 1893 und 1894. — Abh. Senckenb. naturforsch. Ges. 24: 193—288, figs.

Caudell, A. N., 1916. Orthoptera Fam. Locustidae subfam. Mecopodinae. — In: P. Wytsman, Genera
Insectorum 171: 1—32, figs.

Emsley, M. G., 1970. A revision of the Steirodontine katydids (Orthoptera: Tettigoniidae; Phaneropte-
rinae: Steirodontini). — Proc. Acad. nat. Sci. Philad. 122: 125—248, figs.

———, Nickle D. A. & W. Wayne Moss, 1967. The value of the stridulatory file and other characters in
Tettigoniid taxonomy (Orthoptera). — Notul. Nat. 404: 1—9, figs.

———, & Nickle, D. A., 1969. The systematics of Ceraia (Orthoptera: Tettigoniidae: Phaneropterinae).
— Proc. Acad. nat. Sci. Philad. 121: 25—77, figs.

Franssen, C. J. H., 1954. Biologische bestrijding van de sabel-sprinkhaan Sexava nubila St. op de Talau-
de-eilanden. — Ent. Ber. 15: 99—102.

Haan, W. De, 1842—1844. Bijdragen tot de kennis der Orthoptera. — Verh. nat. Gesch. Nederl. overz.
Bezitt. 16 (Zool. 6): 45— 124 (1842); 18 (Zool. 7): 125—164 (1842); 19—20 (Zool. 8—9):
165—228 (1843); 24 (Zool. 10): 229—248, pls. (1844).

Hebard, M., 1922. Studies in Malayan, Melanesian and Australian Tettigoniidae (Orthoptera). — Proc.

Far Acad. nat. Sci. Philad. 74: 121—299, figs.

Huxley, J., 1970. A revision of the genus Catoptropteryx Karsch (Orthoptera: Tettigoniidae). — Bull.
Br. Mus. nat. Hist. (Ent.) 24: 129—170, pl., figs.

Karny, H., 1924. Beiträge zur Malayischen Orthopterenfauna. VIII. Die Mecopodinen des Buitenzor-
ger Museums. — Treubia 5: 137—160, figs.

———, 1926. Fauna Buruana, Tettigoniidae. — Treubia 7: 146—216, figs.

———, 1931. Orthoptera Celebica Sarasiniana. I. Saltatoria. Fam. Tettigoniidae. — Treubia 12
(suppl.): 4—140, figs.

Kästner, A., 1934. Die Sexavae (Mecopodinae) des Stettiner Museums (Orthopteren des Stettiner Mu-
seums 4. Teil). — Stettin. ent. Ztg. 95: 23—53, figs.

Kirby, W. F., 1891. Notes on the orthopterous family Mecopdidae. — Trans. ent. Soc. Lond. 3:
405—412.

———, 1906. A synonymic catalogue of Orthoptera. II: i—viii, 1—562. — London.

Krauss, H. A., 1903. Orthopteren aus Australien und dem Malayischen Archipel, gesammelt von Pro-
fessor Dr. Richard Semon. — Denkschr. med. -naturw. Ges. Jena 8: 743—770, figs.

Leefmans, S., 1927a. A new Sexava species from the island Poat (Celebes). — Treubia 9: 411—412, figs.

———, 1927b. (Locustidae as coconut pests in the Netherlands-Indies and their parasites). — Meded.
Inst. Plantenz. 72: i—v, 1—95, figs.

Linné, C. von, 1758. Systema naturae, 10th ed. 1: 424—433.

Lloyd, J. E. & Gurney, A. B., 1975. Labral stridulation in a katydid (a coconut-infesting “treehopper”’)
(Orthoptera: Tettigoniidae: Mecopodinae). — Ent. News 86: 47—S0, figs.

Moss, W. W., D. A. Nickle & M. G. Emsley, 1970. A polynomial treatment of stridulatory file data
from several species of the katydid genus Ceraia (Orthoptera: Tettigoniidae: Phaneropterinae).
— Notul. Nat. 432: 1—13, figs.

O’Connor, B. A., 1959. The coconut treehopper, Sexava spp., and its parasites in the Madang District.
— Papua New Guin. agric. J. 11: 121—125, figs.

Oudemans, A. C., 1927. Acarologische aantekeningen LX XXVIII. — Ent. Ber. 7: 257—268.

Ragge, D. R., 1955. The wing-venation of the Orthoptera Saltatoria with notes on Dictyopteran wing-
venation: i—vi, 1 —159, figs. — London, Brit. Mus. (Nat. Hist.).

———, 1969. A revision of the African species of Pseudorhynchus Serville (Orthoptera: Tettigoniidae).
— Bull. Br. Mus. nat. Hist. (Ent.) 23: 169—190, figs.

Redtenbacher, J., 1886. Vergleichende Studien über das Flügelgeäder der Insecten. — Annin naturh.
Mus. Wien 1: 153—232, figs.

———, 1892. Monographische Uebersicht der Mecopodiden. — Verh. zool. -bot. Ges. Wien 1892:
183—224, figs.

258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977

Reyne, A., 1960. (Review: J. B. M. van Dinther, Insect pests of cultivated plants in Surinam, Wagenin-
gen, 1960). — Ent. Ber. 20: 231—232.

Sjöstedt, Y., 1933. Orthopterentypen im Naturhistorischen Reichsmuseum zu Stockholm. 4. Tettigonii-
dae. — Ark. Zool. 25A (13): 1—30, figs.

Stal, C., 1873. Orthoptera nova descripsit. — Ofvers. K. VetenskAkad. Forh. Stockh. 30: 39—54.

———, 1874. Recensio Orthopterorum. 2. — Ofvers. K. VetenskAkad. Forh. Stockh. 31: (sep.: 1—8,
1—121).

———, 1877. Orthoptera nova ex Insulis Philippinis. — Ofvers. K. VetenskAkad. Forh. Stockh. 34:
33—58.

Stoll, C., 1813. Représentation exactement colorée d’après nature des Spectres ou Phasmes, des Man-
tes, des Sauterelles, des Grillons, des Criquets et des Blattes, qui se trouvent dans les quatre
parties du monde. Représentation des Sauterelles, des Grillons et des Blattes: 1—28, pl.
la— 13a. — Amsterdam, J. Sepp.

Walker, F., 1870. Catalogue of the specimens of Dermaptera Saltatoria in the collection of the British
Museum. Part 3: 425—604. — London.

Willemse, C., 1933. Résultats scientifiques du voyage aux Indes-Orientales Néerlandaises de
LL.AA.RR. le Prince et la Princesse Léopold de Belgique. Orthoptera II.: fam. Tettigoniidae
and Gryllacridae. — Mém. Mus. r. Hist. nat. Belg., hors série 4(8): 1—15, figs.

———, 1940. On a collection of Indo-australian Tettigoniidae. — Natuurh. Maandbl. 29: 60—64,
69—72, 79—84, figs.

———, 1951. On a collection of Orthoptera from the Caroline Islands from the Bernice P. Bishop Mu-
seum of Honolulu. — Eos Madr., suppl. (1950): 325—362, figs.

———, 1955. Description of some new Orthoptera II. — Natuurh. Maandbl. 44: 36—37, fig.

———, 1957. Notes on Mecopodidae (Orthoptera, Tettigonoidea). — Tijdschr. Ent. 100: 35—42, figs.

———, 1958. On some Tettigonioidea injurious to coconut palms. — Natuurh. Maandbl. 47: 122—125,
figs.

———,, 1961. Tettigonioidea of the Papuan subregion (Orthoptera). I. Mecopodidae. — Pacif. Insects
3: 93—116, figs.

———, 1966. Descriptions of new and redescriptions of lesser known Orthoptera. Part II. — Publties
natuurh. Genoot. Limburg 16: 1—15, figs.

Willemse, F., 1966. List of the types of Orthoptera in the collection of C. Willemse at the Natuurhisto-
risch Museum of Maastricht. — Publties natuurh. Genoot. Limburg 16: 43—73.

Pl. 1 WILLEMSE: Sexavae of Melanesian Subregion 259

4

\
Figs. 1—4. Sexava coriacea (L.): 1, G (Ambon, CW); 2, g (Obi, CW); 3, 9 (Obi, paratype S. grandis,
CW);4, 9 (Sangihe I., ITZ). Figs. 5—6. S. nubila (Stal): 5, 9 (Sorong, CW); 6, 9 (Talaud Is., BMNH).

260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977 Pl. 2

Figs. 7—9. Sexava nubila (Stal): 7, 9 (holotype); 8, ¢ (Mindiptana, CW); 9, & (Beo, Talaud Is., RNH).
Figs. 1O—11. S. karnyi Leefmans: 10, & (Ampana, ITZ); 11, © (Posso, ITZ).

Pl. 3 WILLEMSE: Sexavae of Melanesian Subregion 261

Figs. 12—22. Sexava species: 12—17, male subgenital plate: 12—14, coriacea (L.) (12, Obi, CW; 13,

Ambon, RNH; 14, Halmaheira, CW); 15—16, nubila (Stal) (15, Sorong, CW; 16, Talaud Is., ITZ); 17,

karnyi Leefmans (Ampana, ITZ); 18—22, male cercus: 18—19, coriacea (L.) (18, Ambon, RNH; 19,

Sangihe Is., BMNH); 20—21, nubila (Stal) (20, Mindiptana, CW; 21, Talaud Is., ITZ); 22, karnyi
Leefmans (Ampana, ITZ).

262 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977 Pl. 4

Figs. 23—28. Sexava species, male stridulatory file: 23—25, coriacea (L.) (23, Obi, CW; 24, Halmaheira,
CW; 25, Ambon, RNH); 26—27, nubila (Stal) (26, Koor, CW; 27, Talaud Is., ITZ); 28, karnyi Leefmans
(Ampana, ITZ).

PI.5 WILLEMSE: Sexavae of Melanesian Subregion 263

Fig. 29. Sexava coriacea (L.), stridulatory area of right male fore wing (Obi, CW).

264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977 Pl. 6

31 i 32 ll...

Figs. 30—33. Segestes species: 30, vittaticeps Stal (& lectotype); 31, cornelii sp.n. (& holotype); 32,

unicolor Redtenbacher (3 Koror, Palau, CAS); 33, decoratus Redtenbacher (4 Bubia, CW).

Pl. 7 WILLEMSE: Sexavae of Melanesian Subregion 265

Figs. 34—41. Segestes species: 34, vittaticeps Stal (3 lectotype); 35, unicolor Redtenbacher (& Koror,

Palau, CAS); 36—37, decoratus Redtenbacher (36, 9 holotype; 37, © Murua Agr. Stat, DASF); 38, 41,

stibicki sp.n. (38, & paratype; 41, 9 paratype); 39, brevipennis sp.n. (G holotype); 40, cornelii sp.n. (9
paratype).

|
|
266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977 Pi. 8

Figs. 42—44. Segestes species: 42—43, stibicki sp.n. (42, & holotype; 43, © paratype); 44, brevipennis
sp.n.(¢ holotype).

PI.9 WILLEMSE: Sexavae of Melanesian Subregion 267

Figs. 45—58. Segestes species: 45—47, fastigium of vertex: 45, decoratus Redtenbacher (¢ Casey’s PI.
Popondetta, DASF); 46, cornelii sp.n. (¢ paratype); 47, stibicki sp.n. (Q paratype); 48—52, male
subgenital plate: 48, vittaticeps Stal (lectotype); 49, decoratus Redtenbacher (Siki, DASF); 50, cornelii
sp.n. (paratype); 51, stibicki sp.n. (holotype); 52, brevipennis sp.n. (holotype); 53—58, male cercus: 53,
vittaticeps Stal (lectotype); 54, unicolor Redtenbacher (Koror, Palau, CAS); 55, decoratus Redtenbacher
(Siki, DASF); 56, cornelii sp.n. (paratype); 57, stibicki sp.n. (holotype); 58, brevipennis sp.n. (holotype).

268 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977 PI 10.

(Koror. Palau, CAS); 60, decoratus Redtenbacher (Siki, DASF); 61, cornelii sp.n. (paratype); 62, stibicki

sp.n. (holotype); 63, brevipennis sp.n. (holotype). 64—67, stridulatory area of right male fore wing; 64,

vittaticeps Stal (lectotype); 65, decoratus Redtenbacher (Casey’s PI., Popondetta, DASF); 66, cornelii
sp.n. (paratype); 67, stibicki sp.n. (holotype).

Pl. 11 WILLEMSE: Sexavae of Melanesian Subregion 269

Figs. 68—74. Segestidea species: 68—71, novaeguineae (Brancsik) (68, 3 Kulili Pl., Karkar I., DASF; 69,
Q holotype of Sexava femorata; 70, 9 Hollandia, RNH; 71, 9 Port Moresby, CW); 72—73, rufipalpis
(C. Willemse) (72, & holo-, 73, 9 allotype); 74, uniformis (C. Willemse) (Q holotype).

270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977 PI. 12

Figs. 75—79. Segestidea species: 75—76, uniformis (C. Willemse) (75, & Tulo PI., Manus I., DASF; 76,
® Pak I., DASF); 77—79, gracilis gracilis (C. Willemse) (77, 3 Londolovit, Lihir Is., CW; 78, © Mahur
I., CW; 79, idem).

Pl. 13 WILLEMSE: Sexavae of Melanesian Subregion 271

Figs. 80—84. Segestidea species: 80—82, gracilis simulatrix ssp.n. (80, & holo-, 81, 9 paratype, similar
locality, CW; 82, idem); 83—84, leefmansi (C. Willemse) (83, & holotype; 84, 3 Metakabul PI., New
Ireland, CW).

272 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977 PI. 14

Figs. 85—90. Segestidea species: 85, leefmansi (C. Willemse) (9 Umbukul, New Hanover, DASF);
86—87, acuminata (Kästner) (9 holotype); 88—90, marmorata occidentalis ssp.n. (88—89, ¢ holo-, 90,
© paratype, Hollandia).

PI. 15 WILLEMSE: Sexavae of Melanesian Subregion 273

103 106

Figs. 91—106, Segestidea species: 91—98, male subgenital plate: 91, novaeguineae (Brancsik) (Baliau
Village, CW); 92, rufipalpis (C. Willemse) (paratype); 93, uniformis (C. Willemse) (Bundalis R. C.
Mission, DASF); 94, gracilis gracilis (C. Willemse) (Masahet I., Lihir Is., CW); 95, gracilis simulatrix
ssp.n. (Bainings St. Paul’s, BPBM); 96, leefmansi (C. Willemse) (Teripax PI., Tatau I., CW); 97,
marmorata marmorata Bolivar (Bubia, BPBM); 98, marmorata occidentalis ssp.n. (holotype); 99—106,
male cercus: 99, novaeguineae (Brancsik) (Bubia, CW); 100, rufipalpis (C. Willemse) (paratype); 101,
uniformis (C. Willemse) (Bundalis R. C. Mission, Manus I., DASF); 102, gracilis gracilis (C. Willemse)
(Masahet I, CW); 103, gracilis simulatrix ssp.n. (Bainings St. Paul's, BPBM); 104, leefmansi (C.
Willemse) (Teripax Pl., Tatau I., CW); 105, marmorata marmorata Bolivar (Bubia, BPBM); 106,
marmorata occidentalis ssp.n. (holotype).

274 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977 Pl. 16

Figs. 107—112. Segestidea species, stridulatory file of left male for wing: 107, novaeguineae (Brancsik)
(Kulili pl., Karkar I., DASF); 108, rufipalpis (C. Willemse) (paratype); 109, uniformis (C. Willemse)
(Bundalis R. C. Mission, Manus I., DASF); 110, gracilis gracilis (C. Willemse) (Masahet I., CW); 111,
gracilis simulatrix ssp.n. (Bainings St. Paul’s, BPBM); 112, leefmansi (C. Willemse) (Masahet I., DASF).

PI. 17 WILLEMSE: Sexavae of Melanesian Subregion 275

113

Figs. 113—116. Segestidea species: 113—114, stridulatory file of left male fore wing; 113, marmorata

marmorata Bolivar (Bubia, BPBM); 114, marmorata occidentalis ssp.n. (holotype); 115—116, stridu-

latory area of right male fore wing; 115, novaeguineae (Brancsik) (Kulili Pl., Karkar I., DASF); 116,
rufipalpis (C. Willemse) (paratype).

276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, AFL. 8, 1977 PI. 18 |

Figs. 117—122. Segestidea species, stridulatory area of right male fore wing; 117, uniformis (C.

Willemse) (Bundalis R. C. Mission, Manus I., DASF); 118, gracilis gracilis (C. Willemse) (Matakan PI.,

DASF); 119, gracilis simulatrix ssp.n. (Bainings St. Paul’s, BPBM); 120, leefmansi (C. Willemse)

(Masahet I., DASF); 121, marmorata marmorata Bolivar (Bubia, BPBM); 122, marmorata occidentalis
ssp.n. (holotype).

acuminata
beieri
brevipennis
celebensis
coriacea
cornelii
decoratus
Eumossula
femorata
frater

fuscus

gracilis
gracilis gracilis
gracilis simulatrix
grandis
hanoverana
insulana
karnyi
lanceolata
leefmansi
marmorata
marmorata marmorata
marmorata occidentalis
Moristus
novaeguineae
nubila
princeps
punctipennis
punctipes
rufipalpis
Segestes
Segestidea
Sexava

soror

stibicki
unicolor
uniformis
vittaticeps

WILLEMSE: Sexavae of Melanesian Subregion

INDEX

(Synonyms in italics)

253
229
236
229
220
237
23]
238
242
229
229
248
248
250
220
251
246
226
220
251
254
254
255
218
242
222
242
256
229
245
227
240
218
256
235
230
246
227

277

TIJDSCHRIFT VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

| REGISTER VAN DEEL 120 |

* Een sterretje duidt aan een naam nieuw voor de wetenschap
* An asterisk denotes a name new to science

ARANEAE

alacris I, 3

arctica 18

arcuata 30, 31

arcuatus 30

aspromontis 17, 18, 20

Bathyphantes 25

borealis 24

camtschaticus 7, 38, 45, 47,
[48

concinna 8

concinnella 8

cracens 42

cristatus 3, 5, 6, 24, 40, 47

* drenskyi 6, 22, 47, 48

expunctus 3

falteronensis 17, 20

flavipes 6, 8, 10, 12, 14, 42,
[47

* floriana 6, 8, 28, 47, 48

foliatus 41

gallicus 8, 10

henricae 12, 14

herbicola 6, 7, 16, 42, 47

jacksoni 6, 7, 32, 45, 47, 48

* jacksonoides 6, 7, 33, 34,

[45, 47, 48

leprosus 3, 5, 46

Lepthyphantes | et seq.

Linyphia 8, 12, 15, 17, 36

mengei 6, 8-11, 12, 14, 42, 47

minutus 3

mughi 3

nigriventer 36, 39

nigriventris 6, 7, 36, 39, 45,
[48

pallidiventris 40

pallidus 40

* perseus 5, 11,47, 49

pullatus 13

pygmaea 15

pygmaeus 25

sanfilippoi 18

spiniger 6, 8, 27, 47, 48

suldalensis 42

taczanowskii 19, nota

tarzanowskii 19, nota

tenebricola 6, 7, 8, 12, 13, 30,

[32, 46, 48

tenebricoloides 6, 7, 21, 48,
[49

tenoides 42

tenuis 6, 7, 14, 17, 42, 47, 48,
[49

Theridium 12

tomskica 8

trucidans 19

zebra 42

zebrinus 15, nota

zelatus 42

zibus 42

zimmermanni 6, 8, 12, 14, 15,

[18, 20, 23, 25, 26, 28, 41,
45, 47, 48, 49

COLEOPTERA

* addendum 83, 87

* bicornutum 83, 89

Bolboceras 81

Bolbogonium 77 sqq

* howdeni 87, 97

impressum 82, 92

* insidiosum 83, 95

* pseudopunctatissimum 82,
[90

punctatissimum 87, 96

* scurra 83, 94

triangulum 81, 83, 91

* wiebesi 83, 99

DIPTERA
betulae 153, 154, 164, 165,
[166, 167, 179, 180, 181

279

* brevipalpis 164, 165, 166,
[167, 169, 177, 180, 181
Cecidomyia 153
Clinodiplosis 154
Hormomyia 153
Aligotrophus 153
Semudobia 153 sqq
* skuhravae 164, 165, 166,
[174, 179, 180, 181
* steenisi 164, 165, 166, 167,
[170, 177, 180, 181
* tarda 164, 166, 172, 179,
[180, 181

HETEROPTERA

argentatus 199, 200, 206

Gerris 199 sqq

gibbifer 209, 210

lacustris 199, 208, 210, 212

najas 200

odontogaster 199, 200, 205,
[211

rufoscutellatus 210

thoracicus 199, 200, 207, 210

HYMENOPTERA

Arischnogaster 60
Belonogaster 58
binodis 58
Calligaster 57 nota 3, 58
eilipennis 63, 64
einerascens 57
concinna 58
cristatus 57
cyanopterus 58
Discoelius 57 nota 3
drewseni 63, 64, 69
Eustenogaster 60
fuscus 58

* gracilipes 73

280 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 120, 1977

* Holischnogaster 60, 70
Ischnogaster 55, 56
Ischnogasteroides 56
Liostenogaster 57, 59

* Metischnogaster 59, 60
miniatus 58

montezuma 58
Parischnogaster 58, 60
Plagiolabra 57
prominens 58

spinipes 57

Stenogaster 55, 58, 59
strigosus 58

williamsi 58

Zethus 57, 58

ORTHOPTERA

acuminatus 240, 241, 253
ssp. annulata (C.
[kuekenthali) 146
beieri 229
bimaculata 137
brevipennis 231, 236
caprai 127, 138
carnipes 109, 110, 112
celebensis 229
Chitaura 136
ssp. coerulipes (C. patagiata)
[127
ssp. coerulipes (C. pervittata)
[130, 132
coriacea 219, 220
cornelii 231, 237
Cranae 110, 121 sqq
Cranaella 109 sqq, 127
decoratus 231
emendata 150
Eumossula 240
femorata 211, 240, 242
fuscus 229
* genjam 127, 128, 140
* glabra 127, 128, 147
gracilis 240, 241, 248
grandis 211, 220
haani 136
hanoverana 240, 251
insulana 240, 241, 246
karnyi 211, 226
* kevani 110, 115

kuekenthali 127, 128, 145
lanceolata 211, 220
leefmansi 211, 240, 251
* longipennis 127, 128, 142
luctuosa 127, 128, 146
* manokwari 127, 128, 142
marginata 135, 145
marmorata 240, 241, 254
multicolor 110, 119
nigroreticulata 127, 128, 135
novaeguineae 211, 240, 242
nubila 219, 222
var. obscura (C. rufo-
[femorata) 127, 132, 134
* ssp. occidentalis (S.
[marmorata) 241, 255
Opiptacris 127, 149
Oxya 135
patagiata 123, 127, 128, 130
pervittata 130
pictipennis 127, 149
princeps 240, 242
Pseudocranae 137
punctipes 229, 240, 256
* rammei 110, 118
* rubra 127, 128, 144
rufipalpis 211, 240, 241, 245
rufipes 127, 128, 141
rufofemorata 127, 132
* samarensis 110, 118
Segestes 227 sqq
Segestidea 240 sqq
Sexava 218 sqq
* ssp. simulatria (S. gracilis)
(241, 250
soror 240, 256
stibicki 231, 235
Taeniophora 136
tibialis 127, 128, 132
trivittata 127, 128, 140
tuberculata 110, 112, 116
unicolor 230
uniformis 211, 240, 256
unistrigata(tum) 127, 128,
[135, 151
vittaticeps 227
willemsei 110, 116

PLANTAE

Betula alba 154

Betula albo-sinensis 181
Betula alleghaniensis 180,
[181
Betula alnoides 181
Betula apoiensis 181
Betula x borggraveana 181
Betula celtiberica 181
Betula chichibuensis 181
Betula x coerulea 181
Betula coerulea-grandis 181
Betula corylifolia 181
Betula costata 180, 181
Betula cylindrostachya 181
Betula davurica 179, 181
Betula ermanii 175, 180, 181
Betula forrestii 181
Betula fruticosa 181
Betula glandulifera 181
Betula glandulosa 181
Betula grossa 180, 181
Betula x intermedia 180,
[181
Betula humilis 181
Betula jacquemontii 180, 181
Betula kirghisorum 181
Betula lenta 180, 181
Betula litwinowii 181
Betula maximowicziana 181
Betula medwediewii 181
Betula middendorfii 181
Betula nana 180, 181
Betula nigra 181
Betula occidentalis 153, 170,
[175, 180, 181
Betula papyrifera 153, 170,
[177, 180, 181
Betula pendula 153, 168, 174,
[177, 181
Betula platyphylla 181
Betula populifolia 170, 177,
[181
Betula pubescens 174, 175,
[180, 181
Betula pumila 181
Betula x purpusii 161
Betula raddeana 180, 181
Betula schmidtii 181
Betula tatewakeana 181
Betula turkestanica 181
Betula utilis 181
Cocos nucifera 245, 251
Marasmius 63

UIL

3 2044 093 392 280

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