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HARVARD UNIVERSITY © Library of the

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DEEL 120 1977

TIJDSCHRIFT VOOR ENTOMOLOGIE

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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. Subscription rate: D.Fl. 170.— per year.

Afleveringen 1-2 verschenen 15.11.1977 Afleveringen 3-5 verschenen l.vii.1977 Afleveringen 6-8 verschenen 30.x11. 1977

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

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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.

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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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| 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.

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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